Biological Control of Insect Pests: Southeast Asian Prospects - EcoPort

Biological Control 

of Insect Pests: 

Southeast Asian 

Prospects 

D.F. Waterhouse

Biological Control of 

Insect Pests: 

Southeast Asian Prospects 

D.F. Waterhouse 

(ACIAR Consultant in Plant Protection) 

Australian Centre for International Agricultural Research 

Canberra 

1998

ii 

The Australian Centre for International Agricultural Research (ACIAR) was 

established in June 1982 by an Act of the Australian Parliament. Its primary 

mandate is to help identify agricultural problems in developing countries and to 

commission collaborative research between Australian and developing country 

researchers in fields where Australia has special competence. 

Where trade names are used this constitutes neither endorsement of nor 

discrimination against any product by the Centre. 

ACIAR MONOGRAPH SERIES 

This peer-reviewed series contains the results of original research supported by 

ACIAR, or deemed relevant to ACIARÕs research objectives. The series is 

distributed internationally, with an emphasis on the Third World 

©Australian Centre for International Agricultural Research 

GPO Box 1571, Canberra, ACT 2601. 

Waterhouse, D.F. 1998, Biological Control of Insect Pests: Southeast Asian Prospects. 

ACIAR Monograph No. 51, 548 pp + viii, 1 fig. 16 maps. 

ISBN 1 86320 221 8 

Design and layout by Arawang Communication Group, Canberra 

Cover: Nezara viridula adult, egg rafts and hatching nymphs. 

Printed by Brown Prior Anderson, Melbourne

Contents 

Foreword vii 

1 Abstract 1 

2 Estimation of biological control prospects 2 

3 Introduction 3 

4 Target insect pests 9 

4.1 Agrius convolvuli 

9 

4.2 

4.3 

4.4 

Rating 10 

Origin 10 

Distribution 10 

Biology 10 

Host plants 11 

Damage 11 

Natural enemies 11 

Attempts at classical biological control 14 

Discussion 16 

Anomis flava 

17 

Rating 18 

Origin 18 


Biology 18 


Damage 19 


Attempts at classical biological control 27 

Major natural enemies 31 

Discussion 31 

Aphis craccivora 

33 

Rating 34 

Origin 34 


Biology 34 


Damage 35 


Comments 44 

Aphis gossypii 

45 

Rating 46 

Origin 46 


Biology 46 


Damage 47 

iii

iv 

4.5 

4.6 

4.7 

4.8 

4.9 


Attempts at biological control 60 

The major parasitoid species 70 

An aphid-specific predator 79 

Comments 80 

Cosmopolites sordidus 

85 

Rating 86 

Origin 86 


Biology 87 

Damage 88 




Biology of main natural enemies 103 

Comments 104 

Deanolis sublimbalis 

105 

Synonyms 106 

Rating 106 

Origin 106 


Biology 107 


Damage 109 


Comment 112 

Diaphorina citri 

113 

Rating 114 

Origin 114 


Biology 115 


Damage 116 




Comments 133 

Dysdercus cingulatus 

135 

Rating 136 

Origin 136 


Biology 136 


Damage 137 


Comment 138 

Dysmicoccus brevipes 

141 

Rating 142 

Origin 142 


Taxonomy 142

4.10 

4.11 

4.12 

4.13 

Biology 143 

Hosts 144 

Damage 144 




Comments 156 

Hypothenemus hampei 

157 

Rating 158 

Origin 158 


Biology 159 


Damage 164 



Major parasite species 176 

Comments 180 

Leucinodes orbonalis 

185 

Rating 186 

Origin 186 


Biology 187 


Damage 189 




Comments 194 

Nezara viridula 

197 

Rating 198 

Origin 198 


Biology 199 

Damage 199 


The role of pheromones and other chemical secretions 208 


Biology of the major species 228 

Comments 233 

Ophiomyia phaseoli 

235 

Rating 236 

Origin 236 


Biology 237 


Damage 238 



The more important parasitoids 251 

Comment 255 

v

vi 

4.14 

4.15 

4.16 

Phyllocnistis citrella 

257 

Rating 258 

Origin 258 


Biology 259 


Damage 263 




Comment 285 

Planococcus citri 

287 

Rating 288 

Origin 288 


Biology 289 


Damage 290 



Biology of important natural enemies 311 

Comments 315 

Trichoplusia ni 

317 

Rating 318 

Origin 318 


Biology 318 


Damage 320 


Introductions for biological control of T. ni 

334 

Major parasitoid species 340 

Comment 345 

5 References 349 

6 Index of scientific names of insects 477 

7 General index 531

Foreword 

Since its inception in 1982, ACIAR has been a very strong supporter 

of classical biological control as a key element in the management of 

exotic arthropod and weed pests. When practiced with appropriate 

safeguards, it often provides a sustainable and environmentally 

friendly alternative to the growing use of pesticides, particularly 

when integrated, if necessary, with the use of resistant plant varieties 

and cultural controls. 

Classical biological control has been very successful in regions of 

the world (e.g. Australia, California, New Zealand, Oceania) where a 

large number of the major insect pests and weeds are exotic. This 

situation applies to a far lesser extent to Southeast Asia but, in a 

recent survey commissioned by ACIAR, Waterhouse (1993b) 

identified 40 major arthropod pests that merited evaluation as 

possible targets for biological control. Not all of these (e.g. the 

indigenous fruit flies) are attractive targets, but some at least are. 

The present volume is a companion to Biological Control of 

Weeds: Southeast Asian Prospects (Waterhouse 1994). It 

summarises what is known about the natural enemies (principally the 

parasitoids) of the major exotic insect pests and indicates prospects 

for their biological control. The aim has been to facilitate, for 

countries of the region, the selection of promising individual, or 

collaborative, priority insect pest targets. This should also provide 

donor agencies with a readily accessible overview of the regionÕs 

major exotic insect pest problems and with an evaluation, where 

possible, of prospects for their amelioration by introduction of 

natural enemies. This should assist in the selection, for support, of 

projects that are best suited to their individual terms of reference. 

R. Clements 

Director 

Australian Centre for International 

Agricultural Research 

vii

1 Abstract 

Introduction 1 

Biological control programs have been mounted in some region(s) of the 

world against 13 of the 16 dossier pests and substantial or partial success has 

been achieved in one or more countries for 8. On the basis of available 

information there are good to excellent prospects for reducing, in at least 

some parts of the region, the damage caused by the following: Leucinodes 

orbonalis, 

Nezara viridula, 

Ophiomyia phaseoli and Planococcus citri. 

There are also good reasons for believing that there will prove to be valuable 

natural enemies for the following: Agrius convolvuli, Anomis flava, 

Aphis 

craccivora, 

Aphis gossypii, 

Diaphorina citri, 

Dysmicoccus brevipes, 

Hypothenemus hampei, 

Phyllocnistis citrella and Trichoplusia ni. 

There 

seems to be little prospect for classical biological control of Dysdercus 

cingulatus, too little is known about Deanolis sublimbalis and the prospects 

for control of Cosmopolites sordidus are unclear, although its lack of pest 

status in Myanmar is puzzling.

2 Biological Control of Insect Pests: Southeast Asian Prospects 

2 Estimation of biological control 

prospects 

Insect Rating Family Any Attractiveness 

biological as a target in 

control 

successes 

SE Asia 

Agrius convolvuli 

7 Sphingidae yes medium 

Anomis flava 

10 Noctuidae yes low to medium 


15 Aphididae ? medium 


19 Aphididae yes medium 

Cosmopolites sordidus 13 Curculionidae ? uncertain 

Deanolis sublimbalis 3 Pyralidae no uncertain 


8 Psyllidae yes medium 

Dysdercus cingulatus 11 Pyrrhocoridae no v. low 

Dysmicoccus brevipes 10 Pseudococcidae yes medium 

Hypothenemus hampeii 12 Scolytidae yes medium 

Leucinodes orbonalis 15 Pyralidae no medium to high 


10 Pentatomidae yes high 

Ophiomyia phaseoli 14 Agromyzidae yes high 

Phyllocnistis citrella 16 Phyllocnistidae yes medium 


7 Pseudococcidae yes high 


7 Noctuidae yes medium

3 Introduction 


Waterhouse (1993b) published information, collected from agricultural and 

weed experts in the 10 countries of Southeast Asia, on the distribution and 

importance of their major arthropod pests in agriculture. Ratings were 

supplied on the basis of a very simple system 

+++ very widespread and very important 

++ widespread and important 

+ important only locally 

P present, but not an important pest 

The advantages and limitations of this system were discussed by 

Waterhouse (1993b). Of 160 insect and mite pests nominated as important in 

Southeast Asia, a subset of 47 was rated as particularly so. 

The aim of the present work has been to summarise information relevant 

to the prospects for classical biological control of the most important of 

those of this subset of 47 that are thought to have evolved outside Southeast 

Asia. The assumption is that many of these have been introduced without 

some (sometimes without any) of the natural enemies that help to control 

them where they evolved. The chances are very much lower for arthropod 

pests that evolved in Southeast Asia of introducing effective, sufficiently 

host-specific, organisms from outside the region. On the other hand, there is 

reason to believe that some parasitoids of pests that are thought to have 

arisen in, or adjacent to, the Indian subcontinent may not yet occur 

throughout the eastern region of Southeast Asia and several such pests are 

dealt with. 

In regional considerations of this sort, it is to be expected that not all of 

the top 20, or even the top 10, of any one countryÕs arthropod pests will 

necessarily be included. Indeed, at least some of those omitted might well 

merit the production of additional dossiers if they are of such importance 

locally that a biological control program might be justified. ACIAR would 

be interested to learn of pests that might be considered in this category. 

The summary accounts presented are designed to enable a rapid review 

to be made of (i) the main characteristics of the principal insect pests of 

agriculture that are believed to be exotic to part or all of Southeast Asia, (ii) 

what is known of their enemies, particularly those that have high or 

moderate levels of host specificity and (iii) what the prospects appear to be 

for reducing their pest status by classical biological control.


In most instances four databases (and particularly CABI) were searched 

for relevant information: 

AGRICOLA (Bibliography of Agriculture) 1970+ 

BIOSIS (Biological Abstracts) 1989+ 

CABI (CAB International) 1972+ 

DIALOG (Biological Abstracts) 1969+ 

In addition, in many instances abstracting journals and other published 

sources prior to the above commencement dates were also searched. 

Furthermore, useful information was also obtained from other references 

and from unpublished records. Nevertheless, in many cases the search 

cannot be described as exhaustive. Even more relevant than attempting an 

exhaustive search would be a fresh, detailed, field survey targeted on the pest 

in the region where it is causing problems. This is in order to determine what 

natural enemies are already present and, in particular, whether any of the 

organisms that might be considered for introduction are already present. 

The species dealt with are drawn from tables 4 and 5 of ÔThe Major 

Arthropod Pests and Weeds of Agriculture in Southeast Asia: Distribution, 

Importance and OriginÕ (Waterhouse 1993b). It is quite possible that 

additional arthropod pests rating highly in these tables will prove to be exotic 

to Southeast Asia (or significant parts of it) and, alternatively, that some 

considered to be exotic will, on further evidence, be shown to have evolved 

in the region. The ratings of the pests in the Pacific and Southern China 

included at the beginning of each dossier are based on information in 

Waterhouse (1997) and Li et al. (1997). 

The natural enemies most commonly selected against insect pests in 

modern classical biological practice are specific or relatively specific 

parasitoids. Although predators also clearly play an important role in 

reducing pest numbers (and have achieved considerable successes against 

scale insects and mealybugs) the majority of predators attack a wide 

spectrum of hosts. National authorities responsible for approving the 

introduction of biological control agents are becoming increasingly 

reluctant to do so for natural enemies that may possibly have adverse effects 

on non-target species of environmental significance. For this reason far more 

emphasis has been placed in the dossiers on parasitoids than on predators. 

There appears to be a widespread view that, when biological control 

alone results in a spectacular reduction in pest populations (as it often does) 

it is very worthwhile, but a lesser reduction is of little or no value. Nothing 

can be further from the truth, since far lower levels can have a major impact 

when integrated with other means of pest control. This applies particularly to

Figure 1. 


integration with the use of plant varieties that are partially resistant to the 

pest (Waterhouse 1993a). 

Plant resistance serves to decrease numbers, in particular by lowering 

reproductive rate and slowing growth rate. Resistance can be brought about 

inter alia by alteration of the physical characteristics (e.g. hairiness, cuticle 

thickness) of the plant and/or its chemical composition. If, as usually occurs, 

parasitoids and predators are not affected to an equal extent, an improved 

ratio of natural enemy to the pest will result and the impact of biological 

control will be increased. This was pointed out many years ago (van Emden 

1966; van Emden and Wearing 1965) and is well illustrated by glasshouse 

tests with the aphis Schizaphis graminum on susceptible and resistant barley 

and sorghum varieties and the parasitoid Lysiphlebus testaceipes (Starks et 

al. 1972). If it is assumed, as in the illustrative example in Figure 1, that the 

economic injury level is 100 aphids per plant, then neither the resistant 

variety alone, nor the parasitoid alone will prevent the injury level being 

exceeded, whereas the combination of resistance and parasitoids achieves 

this by a wide margin. As another example, biological control of Myzus 

persicae with Aphidius matricariae was only effective on chrysanthemums 

if the variety involved was partly aphid resistant (Wyatt 1970). 

300 

200 

100 

0 

0 

1 2 

Weeks 

3 4 

Susceptible 

No parasitoid 

Resistant 

Susceptible 

Parasitoid 

present 

Resistant 

Population growth of Schizaphis graminum on susceptible 

and partly resistant barley in the presence and absence of the 

parasitoid Lysiphlebus testiceipes.


Efforts to achieve pest control by high levels of plant resistance alone 

may prove counterproductive if significant energy or other resources are 

diverted by the plant, since they cannot then be used for growth or 

reproduction. Thus, van Emden (1991) quotes data on 31 pigeon pea 

varieties screened at the International Crops Research Institute for the Semi 

Arid Tropics for insect pod damage. These data predicted a 31% yield loss 

for 90% resistance to insects. To accept a loss of this order is surely an 

unacceptable ÔsolutionÕ to the problem, particularly when even a low level of 

natural enemy attack combined with moderate plant resistance is likely to 

achieve a far better yield. However, the interaction of resistance and natural 

enemies may not be a simple one, as pointed out by Wellings and Ward 

(1994) and such interactions urgently deserve further study. Nevertheless, 

the fact remains that, when integrated appropriately with plant resistance and 

other measures, even comparatively low levels of attack by natural enemies 

can lead to disproportionately large improvements in pest control. 

Although the major focus of the dossiers has been on the applicability of 

the information to biological control in Southeast Asia, much has far wider 

applicability. In particular, a great deal is relevant to classical biological 

control in the oceanic Pacific which, until the past few decades, has received 

almost all its important insect pests from Southeast Asia. A brief tabulation 

of the distribution and importance of each pest in the Pacific is, therefore, 

given at the beginning of each dossier. The key to Pacific Country 

abbreviations is: Fr P, French Polynesia; FSM, Federated States of 

Micronesia; Kiri, Kiribati; Mar Is, Marshall Islands; N Cal, New Caledonia; 

PNG, Papua New Guinea; A Sam, American Samoa; Sam, Western Samoa; 

Sol Is, Solomon Islands; Tok, Tokelau; Tong, Tonga; Tuv, Tuvalu; Van, 

Vanuatu; W&F, Wallis and Futuna. The key to Southeast Asian countries is: 

Myan, Myanmar (Burma); Thai, Thailand; Laos; Camb, Cambodia; Viet, 

Vietnam; Msia, Malaysia; Sing, Singapore; Brun, Brunei; Indo, Indonesia; 

Phil, Philippines. 

In any biological control program it is essential that appropriate 

procedures are adopted in relation to the selection of suitably host-specific 

natural enemies, the gaining of approval for introduction and release from 

the national authorities and safe procedures for eliminating unwanted fellow 

travellers. Simple Guidelines for biological control projects in the Pacific 

(Waterhouse 1991) are available from the South Pacific Commission, 

Noumea and FAO has a Draft Code of Conduct for the Import and Release of 

Biological Control Agents (1993) . 

Because there is a considerable lack of uniformity in the names applied 

to many of the insects involved, a separate index is included listing the 

preferred scientific names. These have been used in the text, replacing where


necessary those used by the authors quoted. Where the name of an insect 

used in a publication is no longer preferred by taxonomists, the superseded 

name, x, is shown thus (= x), but this usage is not intended to convey any 

other taxonomic message. Indeed, the superseded name may still be valid, 

but simply not applicable to the particular species referred to by the author. 

I am most grateful for assistance from many colleagues during the 

preparation of this book. It is not possible to name them all, but special 

thanks are due to a number of CSIRO colleagues, in particular to Dr K.R. 

Norris for editorial assistance, Dr M. Carver for valuable advice on the Aphis 

dossiers, J. Prance for bibliographic assistance and to several taxonomists, 

including Dr M. Carver (Hemiptera), Dr P. Cranston (Diptera), E.D. 

Edwards (Lepidoptera), Dr I.D. Naumann (Hymenoptera) and T. Weir 

(Coleoptera). Others who have provided valuable information include D. 

Smith (Queensland Department of Primary Industries), Dr P. Cochereau 

(ORSTOM, Noumea) and Dr C. Klein Koch (Chile). 

Continuing warm support has been provided by Dr P. Ferrar, Research 

Program Coordinator, Crop Sciences, ACIAR, Canberra. 

It is again a pleasure to acknowledge, warmly, the expert assistance of 

Mrs Audra Johnstone in converting my manuscripts into presentable form. 

It would certainly not have been possible to continue with these 

biological control activities long into retirement without the unfailing 

support, encouragement and forbearance of my wife, to whom my very 

special thanks are due.

4 Target insect pests 

4.1 Agrius convolvuli 

India 

20° 

Myanmar 

P Laos 

P 

0° 

20° 

China 

++ 

Thailand 

+ 

Cambodia 

+ 

Vietnam 

++ 

P 

+ Brunei 

Malaysia 

Singapore 

++ 

Indonesia 

Taiwan 

++ 

Philippines 

P 

Australia 

Papua 

New Guinea 

+ 

The moth Agrius convolvuli is widespread in the tropics and subtropics, except for the 

Americas where it does not occur. 

It is an important pest, sporadically, of sweet potato and also attacks several important 

legumes. For most of the time its populations are maintained at subeconomic levels, 

apparently by several trichogrammatid egg parasitoids. These could be considered for 

introduction as biological control agents where they do not already occur. The cause of 

sporadic outbreaks is unknown. 

20° 

0° 

20° 

9


Agrius convolvuli (Linnaeus) 

Rating 

Origin 

Distribution 

Biology 

Lepidoptera: Sphingidae 

sweet potato hawk moth, sweet potato hornworm 

Synonym: Herse convolvuli 

Southeast Asia China Southern and Western Pacific 

++ Viet, Indo ++ + N Cal, PNG 

7 + Thai, Camb 

Msia 

2 

P Myan, Brun P Widespread 

Very widespread in tropical and subtropical areas of the world, except for 

the Americas. 

Southern Europe: Azores, Crete, Malta, Sicily, Yugoslavia. Africa: 

Algeria, Angola, Benin, Burundi, Cape Verde Is, Congo, Egypt, Ethiopia, 

Ghana, Ivory Coast, Kenya, Libya, Madagascar, Madeira, Mali, Mauritius, 

Morocco, Mozambique, Niger, Nigeria, Rwanda, St Helena, Senegal, 

Seychelles, Sierra Leone, Somalia, South Africa, Sudan, Swaziland, 

Tanzania, Togo, Tunisia, Uganda, Upper Volta, Zambia, Zimbabwe. Asia: 

Andaman Is, Bangladesh, Bhutan, Cambodia, China, Christmas Is, Cyprus, 

India, Indonesia, Iran, Iraq, Israel, Japan, Laos, Malaysia, Myanmar, 

Pakistan, Philippines, Saudi Arabia, Singapore, Sri Lanka, Syria, Thailand, 

Turkey, Vietnam. Australasia and Pacific Islands: 

Australia, Cook Is, Fiji, 

Hawaii, Kiribati, Mariana Is, Marquesas Is, New Caledonia, New Zealand, 

Niue, Norfolk Is, Papua New Guinea, Samoa, Solomon Is, Tonga, Tuvalu, 

Vanuatu (CIE Map No 451, 1983). 

The smooth eggs are laid singly on stems and leaves and, in common with 

most other Lepidoptera, A. convolvuli larvae have 5 instars. There is a green, 

a black and a brown form of larvae, which have, at the posterior end, a 

uniformly curved, tapering, smooth dorsal horn. Fully grown larvae attain a 

length of 9 cm. Pupation occurs in earthern cells several centimetres below 

the soil surface. The pupa has a very characteristic proboscis, which is 

enclosed in a looped tube not fused to the body (Kalshoven 1981; Common

Host plants 

Damage 

4.1 


1990). The mean development period at 25¡C in Japan was 21.2 days 

(Setokuchi et al. 1985). In Egypt at 30¡C and 61% RH average 

developmental periods were: larvae 14.4 days, prepupae 1.9 days and pupae 

13 days (Awadallah et al. 1976). The moths often enter houses in the evening 

and, when at rest, resemble pieces of bark. There are at least two generations 

during summer, and winter is passed as a pupa. 

An artificial diet containing powdered sweet potato leaf has been 

developed (Kiguchi and Shimoda 1994). On this at 27¡C and with a day 

length of 16 hours, A. convolvuli larvae moulted to the 5th instar 12 to 14 

days after hatching, pupated at 21 to 26 days and adults emerged at 36 to 41 

days. The 5th instar larvae grew to 8 cm in length and 11 to 12 g in weight 

(Shimoda et al. 1994). Consumption of sweet potato leaves was greatest at 

30¡C, the last instar eating 88% of the total dry weight (5 g) consumed 

(Setokuchi et al. 1986). 

The main commercial host is sweet potato ( Ipomoea batatas), 

but larvae also 

attack other Ipomoea species [e.g. I. pescapreae, 

I. cairica, 

I. indica 

(morning glory) I. hederifolia, 

(Moulds 1981)] and other Convolvulaceae 

[e.g. Merremia dissecta; 

bindweed, Convolvulus arvensis; 

Awadallah et al. 

1976; (Moulds 1981)]. Several pulses are attacked [e.g. wild mung, Vigna 

vexillata (Govindan et al. 1989); moth bean, V. aconitifolia (Bhat et al. 

1990); mung bean, V. radiata and urd bean, V. mungo (Shaw et al. 1989); 

and also Phaseolus spp. (Nagarkatti 1973)]. A strain of moth bean (IPCMO 

131) showed good resistance to attack (Bhat et al. 1990). In Papua New 

Guinea taro is also recorded as a host (Smee 1965). 

A. convolvuli larvae can defoliate sweet potato vines and, even when damage 

is less severe, harvest is delayed, increasing the likelihood of major attack by 

the sweet potato weevil, Cylas formicarius. 

Defoliation of pulses results in 

partial or complete crop failure. 

Natural enemies 

These are shown in Table 4.1.1. 

11

Table 4.1.1 

Natural enemies of Agrius convolvuli 

Species 

DIPTERA 

PHORIDAE 

Country Reference 

Megaselia rufipes 

TACHINIDAE 

Ireland Flemying 1918 

Sturmia dilabida 

Zimbabwe Cuthbertson 1934 

Zygobothria (= Argyrophylax = Sturmia) 

atropivora 

Malaysia 

Zimbabwe 

Zygobothria ciliata (= Sturmia macrophallus) 

Indonesia 

Oman 

Philippines 

HYMENOPTERA 

Corbett & Miller 1933 

Cuthbertson 1934 

Baranoff 1934 

Whitcombe & Erzinclioglu 1989 

Kalshoven 1981 

BRACONIDAE 

Apanteles spp. 

EULOPHIDAE 

China Wu 1983 

species 

ICHNEUMONIDAE 

China Wu 1983 

Amblyteles fuscipennis 

Central Europe Fahringer 1922 

England Morley & Rait-Smith 1933 

Charops bicolor 

China Wu 1983 

Hadrojoppa cognatoria 

Japan Uchida 1924, 1930 

Trogus exaltatorius 

SCELIONIDAE 

England Morley & Rait-Smith 1933 

Telenomus sp. India Nagarkatti 1973 

12 Biological Control of Insect Pests: Southeast Asian Prospects

Table 4.1.1 

(contÕd) Natural enemies of Agrius convolvuli 

Species 

HYMENOPTERA 


TRICHOGRAMMATIDAE 

Trichogramma achaeae 

India Nagarkatti 1973 

Trichogramma agriae 


Trichogramma australicum 


Trichogramma chilonis 

Guam Nafus & Schreiner 1986 

Trichogramma confusum 

India Nagarkatti & Nagaraja 1978 

Trichogramma ?minutum 

Indonesia Leefmans 1929; Kalshoven 1981 

Trichogramma sp. 

FUNGI 

Philippines Shibuya & Yamashita 1936 

Entomophthora sp. Ôgrylli'Õ 

type Japan Kushida et al. 1975 

4.3 


13


Attempts at classical biological control 

CHINA 

GUAM 

INDIA 

A species of Trichogramma, 

possibly T. australicum, (Nagarkatti 1973), has 

been imported on two occasions (Table 4.1.2) to attack the eggs of pest 

Lepidoptera, including Agrius convolvuli, 

but the resulting impact on 

populations of the sweet potato hawk moth is not recorded. 

Table 4.1.2 Attempts at classical biological control of A. convolvuli 

Species From To Year Result Reference 

HYMENOPTERA 


Trichogramma 

?australicum 

Trichogramma 

?australicum 

USA Indonesia before 

1929 

+ Leefmans 1929; 

Nagarkatti 1973 

Philippines Japan 1929 + Shibuya & 

Yamashita 1936; 

Nagarkatti 1973 

In Fujian Province, A. convolvuli larvae were parasitised by Charops bicolor 

(Ichneumonidae), Apanteles spp. (Braconidae) and eulophid wasps (Wu 

1983). 

A. convolvuli is a minor pest of sweet potato on Guam. When sweet potato 

was intercropped with maize, A. convolvuli eggs were parasitised to the 

extent of 70 to 100% by Trichogramma chilonis. 

This parasitoid attacks the 

eggs of a range of sphingids and noctuids, including Ostrinia furnacalis, 

less 

than 20% of whose eggs on maize were parasitised. A. convolvuli colonises 

new sweet potato plantings as soon as cuttings strike and, by the 4th week, 

30 to 60% of its eggs are parasitised. Each large egg produces 13± 

7 

parasitoids, which emerge about 10 days after the host egg is parasitised. It 

was concluded that T. chilonis is a major mortality factor for the sweet potato 

hornworm (Nafus and Schreiner 1986). 

A. convolvuli is an occasional pest of sweet potato, Vigna mungo and Vigna 

radiata. 

Eggs are also laid on the leaves of Colocasia antiquorum and 

Clerodendrum chinense, 

but no significant feeding occurs on these latter 

plants. 

Four species of parasitoid attack the eggs of A. convolvuli near 

Bangalore: Trichogramma australicum, T. achaeae, T. agriae and a species 

of Telenomus (Eulophidae). The abundance of each parasitoid varied with 

the plant species on which the eggs were laid. T. agriae was the commonest 

species in eggs collected on Colocasia, followed by T. achaeae and

4.1 Agrius convolvuli 15 

Telenomus sp., up to a total of 43.6%. T. australicum, followed by 

Telenomus sp., were the main species emerging from eggs on 

Clerodendrum, up to a total of 63.9%. At no time were T. achaeae or 

T. agriae reared from eggs on Clerodendrum. Furthermore, T. australicum 

was reared only twice from Agrius eggs on Colocasia. These results 

highlight the difficulty of reaching decisions on host specificity on the basis 

of laboratory trials in a non-natural environment. 

Up to 49 Trichogramma individuals were reared from a single 

A. convolvuli egg and only in two instances were more than 1 species reared 

from a single egg. These were 7 T. agriae and 4 T. australicum on one 

occasion and 7 T. achaeae and 11 T. australicum in the second. Eggs 

parasitised by Telenomus sp. usually produced 3 to 5 adults and at no time 

did a Trichogramma emerge from the same egg as a Telenomus (Nagarkatti 

1973). Later, an additional parasitoid (Trichogramma confusum) was 

recorded from the eggs of A. convolvuli on Clerodendrum chinense 

(Nagarkatti and Nagaraja 1978). 

Nagarkatti (1973) suggested that the 4 former species might be 

introduced where A. convolvuli is a pest and where they do not already 

occur. 

INDONESIA 

Leefmans (1929) reported the parasitisation of A. convolvuli eggs by 

Trichogramma minutum imported from America. However, Nagarkatti 

(1973) suggests that, from the distribution of T. minutum at that time, it must 

have been T. australicum or some other species of Trichogramma. 

IRELAND 

An adult A. convolvuli produced, soon after capture, many small puparia, 

from which 76 Megaselia rufipes (Diptera, Phoridae) emerged (Flemying 

1918). 

JAPAN 

A species of Trichogramma that parasitises the eggs of Chilo 

suppressalis (= C. simplex) was imported in 1929 from the Philippines. It 

was shown to parasitise also the eggs of A. convolvuli and 10 other species of 

Lepidoptera belonging to several families (Shibuya and Yamashita 1936). 

Nagarkatti (1973) suggests that the species was Trichogramma australicum. 

OMAN 

Adults of the tachinid Zygobothria ciliata emerged from puparia from a 

larva collected on Ipomoea (Whitcombe and Erzinclioglu 1989).


SOUTH AFRICA 

A. convolvuli is a common pest in the eastern part of South Africa. Although 

it is generally not abundant, from time to time large areas of sweet potatoes 

have been almost completely defoliated by it. There are 3 generations a year 

and overwintering occurs as the pupa. Natural enemies include the whitebellied 

stork (Ciconia nigra) which, on occasion, destroys large numbers of 

larvae (Anon. 1927). 

ZIMBABWE 

A. convolvuli larvae on sweet potato were parasitised by the tachinids 

Zygobothria atropivora and Sturmia dilabida, both of which are widely 

distributed in South Africa. The latter parasitoid also attacks larvae of 

Spodoptera exigua (Cuthbertson 1934). 

Discussion 

The majority of the parasitoids recorded as attacking A. convolvuli also 

attack the eggs or larvae of a range of other Lepidoptera living in the same 

environment. Many of these are pest species. Lack of parasitoid specificity is 

a significant advantage when dealing with a strong flying species, such as 

A. convolvuli, which can travel long distances, since the parasitoids are more 

likely to be already present on some other host when adult moths arrive to 

oviposit at a new site. On the other hand, lack of specificity is a disadvantage 

if the non-target species that are attacked include environmentally important 

species, the lowering of whose population density is considered undesirable. 

In the present instance it is clear, from the information outlined earlier 

under India that, whereas the Trichogramma egg parasitoids involved attack 

the eggs of a range of species of Lepidoptera, they do so only when the eggs 

are laid on particular host plants. In this sense they, indeed, display a 

valuable degree of specificity, which should be taken into consideration 

when deciding whether or not to proceed with introductions. 

With these qualifications it is clear that the establishment, in areas where 

they do not already occur, of any or all of 4 Trichogramma species 

(T. achaeae, T. agriae, T. australicum, T. chilonis) is highly likely to lead to 

a reduction to (or at least towards) subeconomic levels in the population of 

A. convolvuli. 

The underlying causes of the sporadic outbreaks of A. convolvuli are 

unknown. Comparatively little work also has been carried out on the 

parasitoids and more detailed studies may well reveal attractive new options 

to pursue.

4.2 Anomis flava 

India 

20° 

Myanmar 

P Laos 

P 

0° 

20° 

China 

++ 

Thailand 

+ 

Cambodia 

+ 

Vietnam 

+++ 

++ 

Malaysia 

Singapore 

Brunei 

+ 

Indonesia 

Taiwan 

Philippines 

Australia 

Papua 

New Guinea 

The noctuid moth Anomis flava occurs widely in Africa, Asia and Oceania, where its larvae 

sporadically, but seriously, damage cotton, okra, kenaf and other Malvaceae: its adults are 

fruit-sucking moths. Its sporadic occurrence suggests that it may be under effective 

biological control for much of the time. 

It is attacked by non-specific predators and by a number of parasitoids. Many of the 

latter attack other Lepidoptera in the same plant environment and appear to be specific to 

larvae in that environment rather than to individual species inhabiting it. 

Further studies are needed to provide information on what the prospects are for 

classical biological control. 

17 

20° 

0° 

20°


Anomis flava (Fabricius) 

Rating 

Origin 

Distribution 

Biology 

Lepidoptera: Noctuidae: Ophiderinae 

cotton semi looper, green semi looper, okra semi looper 

Synonyms: Cosmophila flava, 

Cosmophila indica. 

Cosmophila is 

now regarded as a subgenus of Anomis. 

A. flava does not occur in 

the Americas, where its equivalent is Anomis ( Cosmophila) 

erosa 

(Pearson 1958). Records of A. erosa in the Asian continent should 

be referred to A. flava. 

Southeast Asia Southern China Pacific 

+++ Viet 

10 ++ Msia ++ 

+ Thai, Camb, 

Indo 

P Myan, Laos, Phil present, but not important 

Unclear: could be Africa or Asia. Information available on specific or 

reasonably specific parasitoids possibly favours Africa. 

Africa: 

Central and southern countries, including Angola, Benin, 

Cameroun, Chad, Congo, Ethiopia, Gambia, Ghana, Ivory Coast, Kenya, 

Madagascar, Malawi, Mali, Mauritius, Niger, Nigeria, Senegal, Somalia, 

Sudan, Tanzania, Togo, Uganda, Upper Volta, Zambia, Zimbabwe. Asia: 

Cambodia, China, India, Indonesia, Japan, Korea, Laos, Malaysia, 

Myanmar, Pakistan, Philippines, Sri Lanka, Taiwan, Thailand, Vietnam. 

Australasia and Pacific Islands: 

Australia, Cook Is, Fiji, Mariana Is, 

Marquesas, New Caledonia, Papua New Guinea, Samoa, Solomon Is., 

Tonga, Vanuatu (CIE 1978), Rapa Is, Hawaii (Common 1990). 

Most A. flava eggs are laid on the undersurface of leaves, the young larvae 

are green, those of the last instar measure up to 35 mm in length and bear 

short, lighter-green longitudinal lines and spots. Young larvae skeletonise 

leaves, older larvae consume narrow leaf (roselle) cotton leaves and eat 

irregular holes in broader leaves. Larval survival and growth are greater on

Host plants 

Damage 

4.2 

Anomis flava 

hirsutum than on desi cotton (Sidhu and Dhawan 1979; Kalshoven 1981). 

Pupation occurs in a cocoon spun between leaves. Development times have 

been recorded on a number of occasions (for examples see Table 4.2.1, also 

Schmitz 1968; Yu and Tu 1969), egg to adult taking about 3 weeks or a little 

longer and the number of eggs laid ranging from 158 to 476, depending, in 

part, upon the larval food plant. Groups of larvae normally pass through 5 

moults whereas, when reared singly, up to 22% pass through 6 moults 

(Kirkpatrick 1963; Essien and Odebiyi 1991). There are 5 overlapping 

generations a year in Hunan Province, China, but fewer in some other 

regions (Chen et al. 1991). 

Adults rest in foliage by day and are active in the evening: they are 

attracted to light. 

A. flava is a major, but sporadic, pest of cotton. Larvae also attack many 

other plants, mainly in the family Malvaceae. These include, especially, okra 

( Hibiscus esculentus), 

but also kenaf or Deccan hemp ( H. cannabinus), 

jute 

( H. sabadariffa), 

bele ( H. manihot), 

muskmallow an important medicinal 

plant ( H. abelmoschus), 

shooflower ( H. rosa-sinensis), 

hollyhock ( Althaea 

rosea), 

Arbutilon spp., Sida spp. and Urena spp. (all Malvaceae). However 

they also attack tomato ( Lycopersicon esculentum: 

Solanaceae); cowpea 

( Vigna unguiculata) 

and green gram ( Vigna radiata): 

Fabaceae; sweet 

potato ( Ipomoea batatas): 

Convolvulaceae; as well as melon ( Citrullus 

lanatus), 

Macadamia, 

Ricinus, 

Leea and Amaranthus spp. (Kalshoven 1981; 

Yein and Singh 1981; Croix and Thindwa 1986; Gatoria and Singh 1988; 

Essien and Odebiyi 1991). 

Okra and hemp (kenaf) were the most favoured larval food plants, 

whereas cotton and okra were the most favourable in terms of pupal weight 

and adult fecundity (Rao and Patel 1973). 

When abundant, A. flava larvae are capable of causing serious damage by 

destroying the leaves and buds of cotton and other Malvaceous crops. 

A. flava belongs to the subfamily Ophiderinae of noctuids, the adults of 

which are often fruit-piercing species. In southern China, A. flava is reported 

to be a serious pest of citrus fruit (Li et al. 1997) and, in Korea, A. flava is one 

of a group of fruit-sucking moths observed to damage grapes and pears (Lee 

et al. 1970). 

19

Table 4.2.1 

Average figures (days) for development of Anomis flava 

Stage Rao & Patel 1973 Kalshoven 1981 

Author 

Ferino et al. 1982a Chen et al. 1991 Essien & Odebiyi 1991 

egg development 2 2Ð3 4Ð5 2Ð7 

1st instar larva 3 2.4 

2nd 1.8 1.8 

3rd 1.9 11 12Ð16 2.2 

4th 2 2.2 

5th 3 2.3 

prepupa 1 1.4 

pupa 6.2 6Ð11 7 

eggÐadult 21.1 21 19Ð23 22Ð29 28 

female longevity 31 28 10 4Ð7 19.8 

number of eggs 158 350 492 476 

preÐoviposition 1.25 3 3.3 

oviposition 6 7 12.4 


4.2 

Anomis flava 

A. flava is regarded as of only minor importance in the Pacific, which is 

not surprising since none of its major larval host plants is of much economic 

importance there. 


Those reported in the literature are listed in Table 4.2.2. 

Egg parasitoids are Trichogramma spp., which on occasion can be 

effective: in Mali, 92% parasitisation by Trichogramma sp. was recorded in 

untreated cotton (Pierrard 1970), 12.1% to 15% in the Philippines (Ferino et 

al. 1982a) and 60 to 80% of eggs on cotton were attacked by T. dendrolimi in 

China (Wang et al. 1985, 1988). 

As for larval parasitoids, Apanteles anomidis parasitised 27.5% in China 

(Xie 1984), Aleiodes aligharensi and Aleiodes sp. together 5.2% in Chad 

(Silvie et al. 1989), Charops bicolor 10.2% in China (Xie 1984), Meteorus 

pulchricornis 4.9% in China (Xie 1984), Meteorus sp. 50 to 69.4% in Nepal 

(Neupane 1977) and Winthemia dasyops 2.5% in Chad (Silvie et al. 1989). 

Most other records did not indicate effectiveness or, if they did, it was lower 

than 2.5% parasitisation. 

A. flava pupae are attacked by at least 5 species of Brachymeria 

(Chalcididae). In Madagascar, B. multicolor and B. tibialis parasitised 98% 

of pupae in some fields (Steffan 1958). 

Further details are provided in the country summaries. It is not easy to 

discern a pattern from these although, under some conditions, parasitoids are 

clearly able to have a major impact on A. flava populations. 

Less is known about the effectiveness of predators, although pentatomid, 

carabid, coccinellid, vespid and spider predators have been reported and the 

Indian mynah bird consumed large numbers of larvae when they were 

abundant (Khan 1956). 

Unexplained disappearance of larvae is often attributed to predation, 

although heavy rainfall may sometimes be responsible. 

Bacillus thuringiensis has been recorded in the field from A. flava larvae 

(Yin et al. 1991) and has given promising control on a number of occasions 

(Angelini and Couilloud 1972; Delattre 1973; Anon 1976b; Wilson 1981; 

Chen et al. 1991). 

Both granulosis and polyhedrosis viruses have been recorded in the field 

(Table 4.2.2) and it is possible that virus preparations might be used for 

control. 

21

Table 4.2.2 

Natural enemies of Anomis flava 

Species 

DERMAPTERA 

CARCINOPHORIDAE 


Euborellia pallipes 

HEMIPTERA 

ANTHOCORIDAE 

China Yang 1985a 

Orius minutus 

LYGAEIDAE 

China Wu et al. 1981 

Geocoris sp. 

NABIDAE 


Nabis sinoferus 

PENTATOMIDAE 


Cermatulus nasalis 

Australia Kay & Brown 1991 

Eucanthecona (= Cantheconidia) furcellata 


Oechalia schellembergii 

NEUROPTERA 

CHRYSOPIDAE 

Australia Wilson 1981 

Chrysopa sp. 

DIPTERA 

TACHINIDAE 


?Isyropa 

India Maheswariah & Puttarudriah 1956 

Cadurcia (= Sturmia) auratocaudata Nigeria, Gold Coast Curran 1934 

Camplyocheta (= Elpe) sp. Cameroun Deguine 1991 

Carcelia (= Zenilla) cosmophilae Australia Curran 1934, 1938 

Carcelia kockiana India Sohi 1964 

Carcelia illota (= Zenilla noctuae) Australia Curran 1934, 1938; Kay & Brown 1991 

Cylindromya (= Ocyptera) sp. Senegal Risbec 1950 


Table 4.2.2 (contÕd) Natural enemies of Anomis flava 

Species 

DIPTERA 


TACHINIDAE (contÕd) 

Exorista apicalia India Sohi 1964 

Exorista sorbillans Australia Kay & Brown 1991 

Palexorista inconspicua (= Sturmia bimaculata) Africa Pearson 1958 

Palexorista quadrizonula Africa 

Tanzania 

Crosskey 1970 

Robertson 1973 

Sericophoromyia marshalli South Africa Taylor 1930 

unidentified Chad Silvie et al. 1989 

Philippines Ferino et al. 1982a 

Winthemia dasyops Chad Silvie et al. 1989 

Zygobothria ciliata (= Sturmia macrophallus) India Thompson 1944; Sohi 1964 

HYMENOPTERA 

BRACONIDAE 

Aleiodes aligharensi Chad Silvie et al. 1989; Silvie 1991 

Aleiodes sp. Chad Silvie et al. 1989; Silvie 1991 


Apanteles anomidis China 

Vietnam Xie 1984; Xiong et al. 1994; 

van Lam 1996 

Apanteles spp. India 

Philippines Maheswariah & Puttarudriah 1956; Sohi1964; 

Ferino et al. 1982a 

Apanteles syleptae Chad Silvie et al. 1989; Silvie 1991 

Cotesia (= Apanteles) ruficrus China Woo & Hsiang 1939 

Fiji Lever 1943 


4.2 Anomis flava 23


Species 

HYMENOPTERA 


BRACONIDAE (contÕd) 

Disophrys lutea Tanzania Robertson 1973 

Meteorus pulchricornis (= M. japonicus) China Chu 1934; Xie 1984 

Meteorus sp. nr fragilis Nepal Neupane 1977 

Nyereria sp. Chad Silvie et al. 1989 

Parapanteles sp. Chad Silvie et al. 1989 

Protomicroplitis sp. Chad Silvie et al. 1989 

Sigalphus nigripes 

CHALCIDIDAE 

China He & Chen 1993 

Brachymeria nr aliberti Chad Silvie et al. 1989 

Brachymeria lasus (= B. obscurata) China Chu & Hsia 1935; Woo & Hsiang 1939 


Vietnam van Lam 1996 

Brachymeria madecassa Mauritius Vaissayre 1977 

Brachymeria multicolor Madagascar Steffan 1958 

Brachymeria paolii Tanzania Robertson 1973 

Brachymeria sp. Australia Kay & Brown 1991 

Brachymeria tibialis 

EULOPHIDAE 

Madagascar Steffan 1958 

Euplectrus manilae Philippines Ferino et al. 1982a; Otanes & Butac 1935; 

Otanes 1935 

Tetrastichus howardi (= T. ayyari) India Maheswariah & Puttarudriah 1956, Sohi 1964 



Species 

HYMENOPTERA 


EUMENIDAE 

Delta (= Eumenes) pyriforme Philippines Ferino et al. 1982a 

Eumenes campaniformis 

ICHNEUMONIDAE 

Philippines Ferino et al. 1982a,b 

Charops bicolor China Xie 1984 

Charops sp. Senegal Risbec 1950 

Echthromorpha agrestoria Australia Kay & Brown 1991 

Enicospilus ?samoana Kay & Brown 1991 

Enicospilus dolosus Chad Silvie et al. 1989; Silvie 1991 

Enicospilus sp. Tanzania Robertson 1973 

Mesochorus sp. China Xie 1984 

Metopius sp. Vietnam van Lam 1996 

Xanthopimpla punctata China Woo & Hsiang 1939 

Zacharops narangae 


China Chu 1934; Woo & Hsiang 1939 

Trichogramma chilonis Vietnam Nguyen & Nguyen 1982 

Trichogramma dendrolimi China Wang et al. 1985, 1988 

Trichogramma minutum India Maheswariah & Puttarudriah 1956; Sohi 1964 

Philippines Otanes & Butac 1935 

Trichogramma japonicum Vietnam Nguyen & Nguyen 1982 

Trichogramma sp. Australia Twine & Lloyd 1982 

sp. Mali Pierrard 1970 

spp. Philippines Ferino et al. 1982b 

4.2 Anomis flava 25


Species 

HYMENOPTERA 


VESPIDAE 

Polistes jokahamae China Anon. 1976a 

Polistes sp. China Anon. 1976a 

COLEOPTERA 

CARABIDAE 

Calosoma schayeri Australia Twine & Lloyd 1982 

Lissauchenius venator Cameroun Deguine 1991 

COCCINELLIDAE 

Coccinella septempunctata China Wu et al. 1981 

ARACHNIDA 

Erigonidium graminicolum China Wu et al. 1981 

Misumenops tricuspidatus China Wu et al. 1981 

sp. (Oxyopidae) Philippines Ferino et al. 1982a 

sp. (Thomisidae) 

NEMATODA 


MERMITHIDAE India Mundiwale et al. 1978 

not specified 

BACTERIA 

Chad Silvie 1991 

Bacillus thuringiensis wuhanensis 

VIRUSES 

China Yin et al. 1991 

Granulosis China Yin et al. 1991 

Polyhedrosis Australia Bishop et al. 1978 

Cameroun Delattre 1973 

China Liang et al. 1981 

Mali Atger & Chevalet 1975 

AVES 

Vietnam van Cam et al. 1996 

Acridotheres tristis India Khan 1956 


4.2 Anomis flava 27 

Attempts at classical biological control 

There appear to have been only two attempts (Table 4.2.3). The pentatomid 

bug Podisus maculiventris, a general predator of lepidopterous larvae, was 

introduced from USA (where A. flava does not occur) and liberated in Anhui 

Province, China in 1984. However, it failed to become established, possibly 

due to adverse climatic conditions (Wang and Gong 1987). Trichogramma 

minutum from USA was established, in the Philippines in 1934, but its 

impact is not recorded (Otanes and Butac 1935). 

Table 4.2.3 Attempts at biological control of Anomis flava 

Species 

HEMIPTERA 

PENTATOMIDAE 

From To Year Result Reference 

Podisus maculiventris 

HYMENOPTERA 


USA China 1984 Ð Wang & 

Gong 1987 

Trichogramma minutum USA Philippines 1934 + Otanes & 

Butac 1935 

AUSTRALIA 

Regular releases of Trichogramma nr praetiosum at the rate of 50000 adults/ 

ha were made from November to March on 8 ha of cotton in south eastern 

Queensland. The resulting mean rate of egg parasitisation (49.4%) was 

inadequate to control damage by Helicoverpa spp. and the few eggs of 

A. flava collected were not parasitised, although high levels of parasitisation 

had been reported following the release of the same Trichogramma species 

in northern Western Australia (Twine and Lloyd 1982). Good control on 

cotton in northern New South Wales was obtained with a mixture of Bacillus 

thuringiensis and chlordimeform at a time at which, except for coccinellids, 

natural enemies were scarce, although low numbers of spiders and of the 

pentatomid predator Oechallia schellembergii were present (Wilson 1981). 

A. flava is one of two major pests of kenaf in northern Queensland and 

the Ord Irrigation Area of Western Australia, although natural enemies can 

produce valuable control (Kay and Brown 1991). The tachinids Carcelia 

cosmophilae, C. illota and Exorista sorbillans attack larvae of A. flava and 

other noctuids. Larvae are also attacked by the predator Cermatulus nasalis 

(Pentatomidae) and the parasitoids Brachymeria sp. (Chalcididae), 

Echthromorpha agrestoria and Enicospilus ?samoana (both Ichneumonidae) 

(Curran 1938; Kay and Brown 1991).


CHAD 

Eleven species of parasitoid, 3 species of hyperparasitoid and nematodes 

were reared from A. flava larvae on cotton (Silvie et al. 1989; Silvie 1991). 

Total parasitisation never exceeded 25% and, in 1987, 15.7% of 485 A. flava 

larvae were parasitised. Details are shown in Table 4.2.4. The commonest 

parasitoid was Aleiodes aligharensi which, together with Aleiodes sp. 

accounted for nearly a third of all larvae parasitised. Three hyperparasitoids 

were recorded, about half emerging from species of Aleiodes. The most 

abundant was Mesochorus (= Stictopisthus) africanus (Ichneumonidae) 

followed by Nesolynx phaeosoma (Eulophidae) and Eurytoma syleptae 

(Eurytomidae). All three species were also reared from parasitised larvae of 

other host species (Silvie et al. 1989; Silvie 1991). 

Table 4.2.4 Natural enemies of A. flava larvae on cotton in Chad 

Species % of total larvae 

parasitised 

Primary parasitoids 

DIPTERA 

Other hosts 

TACHINIDAE 

Winthemia dasyops 15.8 Chrysodeixis acuta 

HYMENOPTERA 

BRACONIDAE 

Aleiodes aligharensi 32.9 Earias sp. 

Aleiodes sp. Helicoverpa armigera 

Apanteles syleptae 1.3 Syllepte derogata 

Nyereria sp. 1.3 Syllepte derogata 

Parapanteles sp. 5.3 

Protomicroplitis sp. 

CHALCIDIDAE 

3.9 

Brachymeria nr aliberti 

ICHNEUMONIDAE 

1.3 

Enicospilus dolosus 9.2 

NEMATODA 

Hyperparasitoids 

ICHNEUMONIDAE 

1.3 

Mesochorus (= Stictopisthus) africanus 4.0 

Dead parasitoids 23.7


CHINA 

Since 1970 the cultivation of bluish dogbane (Apocynum venotum) has 

increased greatly in Zhejiang Province, where A. flava is its most important 

pest and 43.7% of semilooper larvae were parasitised. There were two 

braconids, Apanteles anomidis (27.5% parasitisation) and Meteorus 

pulchricornis (4.9%); two ichneumonids, Charops bicolor (10 to 15%) and 

Mesochorus sp. (2.4%); and an unidentified species (0.54%). Mesochorus 

sp. acted as a hyperparasitoid of Apanteles anomidis, but itself parasitised 

about 1% of A. flava larvae (Xie 1984). 

Trichogramma chilonis was reared from the eggs of A. flava on cotton in 

Shanxi (Huo et al. 1988). Inoculative releases of T. dendrolimi in vegetable 

gardens adjacent to cotton fields infested with A. flava resulted in 61 to 81% 

parasitisation of its eggs. By comparison, in pesticide-treated fields nearby, 

parasitisation ranged from 2.5 to 30%. Inundative releases directly in cotton 

fields led to 30 to 80% parasitisation and no additional control measures 

were required (Wang et al. 1985, 1988). 

Polistes jokahamae and Polistes sp. were observed in Hunan Province 

preying on A. flava, the late instar larvae being preferred (Anon. 1976a). 

In Hubei Province, the earwig predator Euborellia pallipes was reported 

to reduce A. flava larval populations by 38 to 65% (Yang 1985a). 

INDIA 

Although A. flava is generally a minor pest, serious outbreaks occur 

sporadically. In Hyderabad State more than 1.5 million acres of cotton were 

affected in one outbreak, with up to 30 larvae per plant consuming 

everything except branches and bolls. Large numbers of the common mynah 

were reported eating the larvae (Khan 1956). In Mysore 70% of A. flava 

larvae on cotton in the field were parasitised by tachinid flies and Apanteles 

spp. In the laboratory, eggs were attacked by Trichogramma minutum and 

pupae by Tetrastichus howardi (Maheswariah and Puttarudiah 1956). 

MADAGASCAR 

The non-specific Brachymeria multicolor was recorded as producing more 

than 95% parasitisation of A. flava larvae on cotton (Steffan 1958; Delattre 

1973). B. madecassa was also credited with 50 to 90% parasitisation of 

larvae in 1956 and 1957 (Vaissayre 1977). 

NEPAL 

The most important parasitoid of A. flava larvae, Meteorus sp. nr fragilis 

(Braconidae), was responsible for 50 and 69.4% parasitisation in 1973 and 

1974 respectively. There were no pupal parasitoids (Neupane 1977).


PHILIPPINES 

High temperatures inhibited and moderate rainfall favoured high 

populations of A. flava on seed cotton, yield being significantly reduced only 

at densities of 6 to 8 larvae or greater per plant or at damage rates involving 

at least 60% defoliation. Ten species of natural enemies were recorded, the 

most important being 2 Trichogramma egg parasitoids, a eulophid larval 

parasitoid, a larval and pupal predator (Delta (= Eumenes) pyriforme) and a 

pupal parasitoid (Brachymeria lasus). Larval disappearance was attributed 

to predators, including Eumenes campaniformis and 2 species of spiders. 

Egg and pupal parasitisation were generally high during the wet season, 

whereas larval and pupal predation were higher in the dry season. The major 

mortality occurred during the larval stage, followed by pupal mortality, with 

egg mortality being least important. Larval disappearance, suspected to be 

due to predation, was more important than parasitisation (Ferino et al. 

1982a,b). 

TAIWAN 

A. flava larvae feed on the leaves and buds of kenaf and heavy infestation 

reduces top growth. There are 3 generations a year, of which the 3rd occurs 

in July and is the most injurious. In Taiwan the main hosts are cotton and 

kenaf, although other Malvaceae are attacked (Yu and Tu 1969). The 

biology of Eucanthecona furcellata, a pentatomid predator of A. flava 

larvae, was studied by Chu and Chu (1975). 

TANZANIA 

Four species of parasitoid were reared from A. flava larvae collected from 

cotton and kenaf. In 1963, 7% and, in 1964, 13.8% of larvae were 

parasitised. The species involved were the tachinid fly Palexorista 

quadrizonula, which produced 1 to 5 puparia from each parasitised larva and 

had an average pupal period of 8 days; the ichneumonid Enicospilus sp. 

producing 1 pupa, with an average pupal period of 13 days; and, of lesser 

importance, the braconid Disophrys lutea (1 pupa, 5 days) and the chalcid 

Brachymeria paolii (1 pupa, 11 days). Palexorista quadrizonula was also 

reared from Spodoptera exigua, S. littoralis and Xanthodes graellsii (all 

Noctuidae); Enicospilus sp. from Helicoverpa armigera; and Disophrys 

lutea from Earias biplaga, Spodoptera exigua and S. littoralis (Robertson 

1973). 

VIETNAM 

Control of A. flava is particularly good in some years due to two naturally 

occurring egg parasitoids, Trichogramma chilonis and T. japonicum, 93% 

parasitisation of eggs being reported (Nguyen and Nguyen 1982; Nguyen 

1986).

Major natural enemies 


Apanteles anomidis Hym.: Braconidae 

A. anomidis is an important endoparasite of A. flava in China. It has one 

generation a year. A mated female lays an average of 109 eggs and prefers to 

lay in 1st to 3rd instar host larvae. Adults fed on 10% aqueous sugar solution 

lived about 1.5 days at 29¡C (Xiong et al. 1994). An average of 13.7 pupae of 

A. anomidis were obtained from each parasitised A. flava larva (Xie 1984). 

Palexorista quadrizonula Dip.: Tachinidae 

This parasitoid was the most important of 4 species attacking A. flava in 

Tanzania. It is widespread in Africa south of the Sahara and occurs also in 

the Seychelles and St Helena. It attacks a range of lepidopterous larvae, 

especially species belonging to the Noctuidae, but also to the Arctiidae, 

Geometridae, Pyralidae and Tortricidae. In A. flava it produces 1 to 5 

puparia from each larva, with an average developmental period of 7.9 days 

(Crosskey 1970; Robertson 1973). 

Discussion 

Many natural enemies of A. flava have been reported, although there have 

been few studies detailed enough to indicate their true effectiveness. Most of 

the parasitoids are unlikely to be specific to A. flava, but to attack also other 

lepidopterous larvae feeding on the same host plants. Most of these other 

hosts are themselves pest species, whose abundance it is desirable to lower. 

Specificity in these circumstances is rather to lepidopterous larvae in a 

particular habitat and the parasitoids may thus be sufficiently restricted in 

their attack on non-target species to be seriously considered as agents for 

classical biological control. Indeed, for a sporadic pest such as A. flava, it is 

highly desirable that there should be readily available a reservoir of natural 

enemies present continuously, so as to be in place when populations of 

A. flava start to increase. 

The reasons for sporadic outbreaks have not been identified, although 

Brader (1966) suggested that it might well be due to the application of 

insecticides resulting in the death of natural enemies.

Table 3 

Order No. 

of +s 

26 

1. 41 

2. 35 

3. 34 

4. 32 

5. 31 

6. 30 

7. = 29 

7. = 29 

9. 27 

10. 27 

11. 26 

12. 25 

13. 25 

14. = 24 

14. = 24 

16. 24 

17. 22 

18. 22 

19. 21 

20. 20 

21. = 18 

21. = 18 

23. 17 

24. 17 

25. 17 

26. 16 

27. 16 

28. 15 

29. 15 

30. 15 

a 

Walker 1993. 

Aggregated ratings of the major invertebrate pests of agriculture in the region. 


Pest and 

+ scores 

30 and over 

No. times 

in top 10 

Dossier 

available? 

Any biological 

control successes? 

Attractiveness 

as a target 

Bactrocera spp. 13 + + + 


Spodoptera litura 


Cylas formicarius 

Plutella xylostella 

25–29 

Crocidolomia pavonana 

4 + + + + 

4 + – – 

6 + + + + + 

7 + – – 

9 + + + + + + 

4 + – + 

Liriomyza spp. 4 + + + + 

Othreis fullonia 

Helicoverpa armigera 

Pentalonia nigronervosa 

8 + + + + + + 

4 + + + 

4 + – + 

Epilachna spp. 4 + + + + 

Aulacophora spp. 2 + – – 

20–24 

Nacoleia octasema 

Maruca vittrata 

Polyphagotarsonemus 

latus 

Agonoxena argaula 

Brontispa longissima 

Tarophagus proserpina 

Aleurodicus dispersus 

15–19 

Phyllocnistis citrella 

Unaspis citri 

3 + + + + 

3 + – + 

1 + – – 

5 + + + + + 

4 + + + + + + 

3 + – + + + + 

3 + + + + + + 

2 + + + + + 

2 – + + + + 

Papuana spp. 5 + – + 

Adoretus versutus 


Euscepes postfasciatus 

Halticus tibialis 

Oryctes rhinocerus 

Thrips palmi 

Coccus viridis 

4 + – – 

– + + + + 

3 – – – 

2 – – – 

3 + + + + + + 

3 a (+) 

– – 

1 – ? + + 

(cont’d over)

Table 3 

(cont’d) 

Order No. 

of +s 

31. 14 

32. 14 

32. = 14 

32. = 14 

35. 13 

36. = 12 

36. = 12 

38. 12 

39. 11 

40. 11 

41. 10 

42. 10 

43. = 10 

43. = 10 

43. = 10 

46. 9 

b 

De Barro 1995. 

Aggregated ratings of the major invertebrate pests of agriculture in the region. 

10–14 

Achatina fulica 

Phyllocoptrupa oleivora 

Hellula spp. 


Aspidiotus destructor 

Graeffea crouanii 

Planococcus pacificus 

Earias vittella 

Pest and 

+ scores 


Tetranycus lambi 

Bemisia tabaci 

Ceroplastes rubens 

Hippotion celerio 

Rhabdoscelus obscurus 

Tetranycus marianae 

Still invading 

Bemisia argentifolii 

No. times 

in top 10 

Dossier 

available? 

2 + + + + + + 

– – – – 

– + – – 

– + + + + + + 

2 + + + + + + 

2 + + + + + 

2 – – + 

1 – – – 

1 + + + + 

– – – – 

3 b (+) 

+ + 

1 – + + + + 

– – – – 

– – + + + 

– – – – 

3 b (+) 

Any biological 

control successes? 

Attractiveness 

as a target 

+ + 

The Major Invertebrate Pests and Weeds of Agriculture and Plantation Forestry in the Southern and Western Pacific 

27

4.4 Aphis gossypii 

India 

Myanmar 

++ 

20° 

Laos 

+ 

0° 

20° 

China 

+++ 

Thailand 

+++ 

Cambodia 

++ 

Vietnam 

++ 

+ 

++ Brunei 

Malaysia 

+ 

Singapore 

++ 

Indonesia 

Taiwan 

++ 

+++ 

Philippines 

Australia 

Papua 

New Guinea 

+ 

The comments under the map of Aphis craccivora apply also to A. gossypii. 

45 

20° 

0° 

20°


4.4 Aphis gossypii Glover 

Rating 

Origin 

Distribution 

Biology 

Hemiptera: Aphididae 

cotton aphid, melon aphid 


+++ Thai, Phil +++ +++ Fiji, Guam, Tong, Van 

19 ++ Myan, Camb, Viet, 

32 ++ Cook Is, FSM, Fr P, Kiri, 

Msia, Indo 

Niue, Sam, Tuv 

+ Laos, Sing, Brun + N Cal, PNG, A Sam, Sol 

Is, Tok, W & F 

P P Tuv, Van 

Unclear. Starù (1967a) suggests Ôprobably steppe areas of the Palaearctic 

regionÕ, possibly inferring southeastern Europe and adjoining regions. The 

taxonomic status of A. gossypii is complex and there are a number of 

biotypes. 

A. gossypii is now very widespread throughout warm temperate, subtropical 

and tropical regions of the world. 

The cotton aphid varies greatly in colour, usually from light green or dark 

green to almost black but, for older, overcrowded larvae (nymphs) and at 

high temperatures it is yellow to almost white and the aphids are smaller than 

on young growth. Wingless females (apterae) vary from 0.9 to 1.8 mm in 

length and winged females (alatae) 1.1 to 1.8 mm. 

In Europe, there is no sexual reproduction, but there is in East Africa, 

USA, China and Japan. However, in Japan, there are also parthenogenetic 

overwintering populations (Komazaki 1993). The young generally moult 4 

times (range 3 to 5). Their rate of development is influenced by the host 

plant, cotton being superior to squash. On cotton and squash it takes an 

average of 4.5 and 6.7 days respectively to the adult stage at about 27¡C: 

there is then a period of about 2 days before nymphs are produced. In this 

series of experiments females on cotton produced an average of 27 nymphs 

(range 9 to 43), whereas those on squash produced an average of 14 (range 2

Host plants 

Damage 

4.4 


to 35) (Khalifa and Sharaf 1964). Life history data on cucumber is provided 

by van Steenis and El Khawass (1995). 

In U.K., apterae lived 16 days and each produced about 40 offspring. In 

founding colonies without competition, the 40 progeny could be produced in 

about 7 days and the total population increased about 10 fold each 

subsequent week. The rate was reduced as crowding occurred and only then 

was it possible for the rate of parasitoid increase to exceed that of the aphid 

(Hussey and Bravenboer 1971). 

A. gossypii is widely polyphagous. Cotton, in particular, can carry very 

heavy infestations, as also can various cucurbits (e.g. cucumber, squash, 

watermelon). In many parts of the world it is one of the most serious of the 

aphids on citrus. A. gossypii also infests beans, egg plant, guava, mango, 

okra, paprika, potato, taro and numerous ornamentals. In Central and South 

America it also damages coffee and cocoa. 

As its common and scientific names imply, cotton can be seriously damaged 

by A. gossypii. 

It can be a major problem and even cause death of the plant at 

early stages of growth; and a further serious attack may occur when the plant 

is near maturation and copious production of honeydew can contaminate the 

cotton lint. 

On all of its many hosts, severely attacked leaves curl and young growth 

is stunted. As populations build up, the upper surface of leaves and fruit 

becomes contaminated with honeydew, leading to growth of sooty moulds, 

which is unsightly and interferes with photosynthesis. 

For many crops, virus transmission is far more important than direct 

damage, since even small numbers of migrating aphids can cause serious 

problems, whereas even large colonies may cause only moderate leaf 

deformation (Barbagallo and Patti 1983). Although, formerly, it was not an 

effective vector of citrus tristeza virus, it has now become a dangerous one in 

USA and Israel. Both adults and nymphs can transmit the virus (Komazaki 

1993). A. gossypii is also an important vector of a very wide range of other 

plant viruses. 

47



Two groups of hymenopterous parasitoids attack (but are restricted to) 

aphids, a larger one consisting of species belonging to the family Aphidiidae 

and a smaller group belonging to the family Aphelinidae. Both groups occur 

worldwide as solitary endoparasitoids. Although many of the species are 

recorded as having an extensive host range, there is almost always a 

significant degree of host restriction. Hosts are frequently some (but not all) 

of the species in a particular aphid genus or several closely related genera. 

There is good evidence that there are biotypes within some parasitoid 

species, since populations from some hosts or some areas parasitise a 

narrower range of hosts than the species as a whole. There may also be 

differences between biotypes in their preference for the host aphid when 

feeding on a particular host plant or in a particular habitat. When a parasitoid 

is abundant on a preferred host it may occasionally attack a nearby nonpreferred 

host, as with Diaeretiella rapae, 

a major parasitoid of the cabbage 

aphid Brevicoryne brassicae, 

which has occasionally been recorded from 

both A. craccivora and A. gossypii, 

but for which it exhibits a low preference 

(Dhiman et al. 1983). There are some species (or biotypes of species) that 

have been found capable of generally causing high levels of parasitisation of 

A. craccivora. 

Those selected by Starù (1967a, b) are shown in bold italics in 

table 4.4.1 and might be considered first as potential species for biological 

control introductions to areas where they do not already occur. Valuable 

reviews of the effectiveness of aphid parasitoids are provided by Carver 

(1989), Hagen and van den Bosch (1968) and Hughes (1989). 

Although many coccinellids, syrphids, chrysopids, hemerobiids and a 

few predator species from other insect families attack aphids, their impact in 

regulating populations is generally regarded as disappointing, although they 

must certainly at times limit economic damage. The efficiency of a predator 

depends upon its searching ability and effectiveness in capturing prey. The 

numbers of predators seem to be greatest when aphid numbers are already 

declining after a peak in abundance and, thus, their apparently great impact 

at that time may actually have little significance in population regulation. 

Predators can increase rapidly in numbers only after their prey has become 

sufficiently abundant, so there is an important time lag between prey and 

predator numbers (Hemptinne and Dixon 1991). 

Coccinellids have been used successfully for the biological control of 

several, relatively sessile, coccid pests, whereas results have generally been 

poor against aphids. One of the reasons is that adult coccinellids and their 

larvae are poor at capturing other than first instar aphids (Dixon 1989). 

Indeed, the survival of newly-hatched beetle larvae is very dependent upon

4.4 


the abundance of young aphids, so there is a need for coccinellids to lay eggs 

very early in the development of aphid colonies. Oviposition late in aphid 

population development may result in older larvae starving from lack of 

food and the comparatively poor searching ability of coccinellids for low 

aphid populations aggravates the situation (Hemptinne and Dixon 1991). 

Another reason is that coccinellids disperse when prey populations fall to 

low levels. 

Adults of most aphidophagous syrphids are attracted to, and lay their 

eggs in or close to, large aphid colonies, the number of eggs deposited 

increasing as aphid density increases (Chandler 1967). Syrphid larvae also 

generally become abundant when the aphid colony is already declining. The 

larvae of the aphidophagous cecidomyiid Aphidoletes aphidimyza appear to 

have adequate host specificity to be acceptable for biological control 

introductions. The species has a high degree of density dependence, kills 

more aphids than it consumes and is less affected than many other predators 

by insecticides (Meadow et al. 1985). 

Chrysopids and hemerobiids are more effective than many other 

predators at capturing prey and are likely to be more efficient predators at 

low aphid densities. 

A particular problem with most predators is that they are highly 

polyphagous. They will almost always attack a very wide range of nontarget 

insects, some of which are likely to be of environmental concern. 

Regulatory authorities responsible for approving import permits to a country 

are becoming increasingly reluctant to do so, unless an adequate degree of 

specificity has been demonstrated and this is occasionally possible. 

For the above reasons, no attempt has been made to assemble lists of the 

many generalist predators recorded as attacking (or probably attacking) 

A. craccivora or A. gossypii in the field, although a few facts about their 

activities are recorded in the segments dealing with individual countries in 

order to provide an entry into the literature. Abstracts of many additional 

papers are available in CABIÕs Review of Agricultural Entomology and its 

predecessor Review of Applied Entomology, Series A. The major 

parasitoids of A. gossypii are listed in Table 4.4.1. 

49

Table 4.4.1 

Parasitoids of Aphis gossypii 


HYMENOPTERA 

APHELINIDAE 

Aphelinus abdominalis 

China Shi 1980 

(= Aphelinus sp. nr flavipes) 

Guam 

Fulmek 1956 

India 

Ramaseshiah & Dharmadhikari 1969 

(= Aphelinus flavipes) 

Shanghai Shi 1980 

Aphelinus asychis 

Italy Ferrari & Nicoli 1994 

Aphelinus chaoniae 

Italy Ferrari & Nicoli 1994 

Aphelinus gossypii (= Aphelinus kashmiriensis) 

Australia 

Carver et al. 1993 

Cook Is 

Walker & Deitz 1979 

Hawaii 

Timberlake 1924; Yoshimoto 1965 

India 

Bhat 1987 

Japan 

Takada & Tokomaku 1996 

Tonga 

Carver et al. 1993; 

Stechmann & Všlkl 1988 

Aphelinus humilis 

Australia M. Carver pers. comm. 

Aphelinus mali 

China 

Shi 1985 

India 


Senegal 

Risbec 1951; Fulmek 1956 

Shanghai 

Shi 1980 

Taiwan 

Takada 1992 

Trinidad 

Bennett 1985 

Aphelinus paramali 

Israel Zehavi & Rosen 1988 

Aphelinus semiflavus 

USA Hartley 1922; Spencer 1926; 

Oatman et al. 1983b; Trumble & Oatman 1984 

Aphelinus varipes (= A. nigritus) 

Transcaucusus 

Fulmek 1956 

USA 

Wharton 1983 


Table 4.4.1 (contÕd) 

HYMENOPTERA 

APHELINIDAE (contÕd) 

Aphelinus sp. 

2 ´ spp. 

APHIDIIDAE 

* Aphidius colemani 

Aphidius ervi 

Aphidius floridaensis 



Colombia 

India 

Japan 

Angola 

Argentina 

Australia 

Chile 

China 

Egypt 

India 

Japan 

Kenya 

Mozambique 

Pakistan 

RŽunion 

Tonga 

Uruguay 

Venezuela 

Morocco 

Uzbekistan 

USA, West Indies Starù 1967a,b 

Ramirez & Zuluaga 1995 


Takada 1992 

Starù & van Harten 1972 

Starù 1967a, 1972 

Carver & Starù 1974; Room & Wardhaugh 1977 

Prado 1991, Starù 1975 

Xi & Zhu 1984 

Selim et al. 1987 

Starù 1972; Agarwala et al. 1981 

Starù 1967a 

Starù & Schmutterer 1973 

Starù & van Harten 1972 

Starù 1975 

Starù 1975 

Carver et al. 1993 

Starù 1975 

Cermeli 1989 

Fulmek 1956 

Starù 1979 

4.4 


51


HYMENOPTERA 

APHIDIIDAE (contÕd) 

* Aphidius gifuensis 

China 

Hawaii 

India 

Japan 

Korea 

Taiwan 

Aphidius urticae (= Aphidius lonicerae) 

Bšrner et al. 1957 

Aphidius matricariae (= Aphidius phorodontis) 

Brazil 

Canada 

Chile 

Germany 

India 

Italy 

Lebanon 

Pakistan 

Peru 

Tunisia 

USA 

Aphidius picipes 

Aphidius similis 

Aphidius sonchi 

Aphidius uzbekistanicus 



Shi 1980; Takada 1992 

Mackauer & Starù 1967; Takada 1968 

Raychaudhuri 1990 

Mackauer & Starù 1967; 

Takada 1968, 1992 

Mackauer & Starù 1967; Takada 1992 

Mackauer & Starù 1967 

Starù 1967a 

Starù 1967a 

Prado 1991 

Mackauer 1962b 

Agarwala et al. 1981; Agarwala 1983 

Starù 1976 

Tremblay et al. 1985 

Starù & Ghosh 1983 

Starù 1967a 

Halima-Kamel 1993 

Starù 1967a 


China Li & Wen 1988; Xi & Zhu 1984 

India Agarwala et al. 1981 


India Raychaudhuri 1990; Takada 1992 



HYMENOPTERA 


Aphidius spp. India Agarwala et al. 1981 

Cristicaudus nepalensis 

India Raychaudhuri 1990; Takada 1992 

Diaeretiella rapae 

Ephedrus nacheri 

* Ephedrus persicae 

Ephedrus plagiator 

* Lipolexis gracilis 



India, Japan 

Tunisia 

USA 

Uzbekistan 

Agarwala et al. 1981; Takada 1992 


Starù 1967a 

Starù 1979 

China, Japan, Europe Takada 1968, 1992 

India 

Iraq 

Korea 

Lebanon 

Taiwan 

USA 

USSR 

India 

Uzbekistan 

Japan, 

Korea, Taiwan 

USSR 

China 

Europe 

Hong Kong 

India 

Japan 

Lebanon 

Shanghai 

Taiwan 

Agarwala et al. 1981, Takada 1992 

Al-Azawi 1970 

Takada 1972b; Chou 1981; Paik 1975 


Chou 1981b 

Schlinger & Hall 1960 

Starù 1970 


Starù 1979 

Takada 1992 

Paik 1975; Chou 1981b 

Starù 1970 

Shi 1980; Xi & Zhu 1984; Takada 1992, 

Starù 1970 

Takada 1992 


Takada 1992 


Shi 1980 

Chou 1981b, Takada 1992 

4.4 


53

Table 4.4.1 (contÕd) Parasitoids of Aphis gossypii 

HYMENOPTERA 


Lipolexis scutellaris (= Lipolexis pseudoscutellaris) Hong Kong 

India 

Malaysia 

Philippines 

Vietnam 

Lysaphidus schimitscheki India Raychaudhuri 1990 

*Lysiphlebia japonica China, Japan 

Korea, Taiwan 


Takada 1992 

Starù & Ghosh 1975; Agarwala et al. 1981; Pramanik & 

Raychaudhuri 1984; Raychaudhuri 1990; Takada 1992 

Ng & Starù 1986; Takada 1992; 

V.J. Calilung pers. comm. 1995 

Starù & Zelenù 1983 

Takada 1968, 1992; Paik 1975; Chou 1981; Tian et al. 

1981; Xi & Zhu 1984 

Lysiphlebia mirzai Vietnam Starù & Zelenù 1983; Takada 1992 

*Lysiphlebus fabarum 

Algeria, Bulgaria, Corsica, Starù et al. 1975; Starù 1976 

(= Lysiphlebus ambiguus 

Israel, Italy 

= Lysiphlebus cardui 

Egypt 

Selim et al. 1987 

= Lysiphlebus confusus) 

Europe 

Starù 1970 

Greece 

Santas 1978 

Iraq 

Al-Azawi 1966 

Japan 

Takada 1992 

Lebanon 


Morocco 

Starù 1967a 

Pakistan 

Hamid et al. 1977 

Tunisia 


USA 

Starù 1967a 

USSR 

Starù 1967a; Lyashova 1992 

Uzbekistan 

Starù 1979 

Lysiphlebus shaanxiensis China Chou & Xiang 1982 



HYMENOPTERA 


*Lysiphlebus testaceipes 

Chile 

Colombia 

Cuba 

France 

Guadeloupe 

Haiti 

Hawaii 

Italy 

Mexico 

Spain 

Portugal 

Trinidad 

USA 

Venezuela 

West Indies 

Lysiphlebus sp. Argentina 

Colombia 

Hawaii 

India 

Pakistan 

Portugal 

USA 


Prado 1991 

Fulmek 1956; Vergara & Galeano 1994 

Starù 1967b, 1981 

Starù et al. 1988a,b 

Starù et al. 1987 

Fulmek 1956 

Starù 1967a 

Tremblay & Barbagallo 1982 

Starù & Remaudi re 1982 

Starù et al. 1988a,b; 

Costa & Starù 1988; Starù et al. 1988c 

Bennett 1985 

Spencer 1926; Schlinger & Hall 1960; Starù 1970; 

Oatman et al. 1983b; Trumble & Oatman 1984 

Cermeli 1989 

Starù 1967b 

Fulmek 1956 

Ramirez & Zuluaga 1995 

Fulmek 1956 

Agarwala et al. 1981 

Mohyuddin & Anwar 1972, 1973 

Boelpaepe et al. 1992 

Fulmek 1956 

Praon abjectum India Raychaudhuri 1990; Takada 1992 

Praon absinthii India Agarwala et al. 1981 

Praon exsoletum Uzbekistan Starù 1979 

4.4 Aphis gossypii 55



HYMENOPTERA 


Praon myzophagum India Agarwala et al. 1981 

Praon volucre 

Lebanon 


Tajikistan, Uzbekistan Starù 1979 

Praon sp. 

India 


Uzbekistan 

Starù 1979 

Toxares macrosiphophagum India Raychaudhuri 1990; Takada 1992 

Trioxys acalephae 

China 

Takada 1992, 

India 


Trioxys angelicae 

Corsica, Greece, Iraq, Italy, Starù 1976; Santas 1978 

Israel, Morocco 

Al-Azawi 1970 

Israel 

Rosen 1967b 

Lebanon 

Hussein & Kawar 1984; Tremblay et al. 1985 

Tunisia 


Trioxys asiaticus Iran, Tajikistan, Uzbekistan Starù 1979 

Trioxys auctus 

USSR 

Uzbekistan 

Fulmek 1956; Mackauer & Starù 1967 

Starù 1979 

Trioxys basicurvus India Agarwala et al. 1981; Raychaudhuri 1990; Takada 1992 

Trioxys communis China 

Japan, Korea, Taiwan 

Philippines 

Shi 1980, 1985 

Paik 1976; Chou 1981a; Lu & Lee 1987; Takada 1992 

V.J. Calilung pers. comm. 1995 

Trioxys complanatus Uzbekistan Starù 1979 

Trioxys equatus India Samanta et al. 1985; Raychaudhuri 1990 



HYMENOPTERA 


*Trioxys indicus China 

Tian et al. 1981, Xi & Zhu 1984 

India 

Starù & Ghosh 1975; Agarwala et al. 1981; Agarwala 

1983,1988; Raychaudhuri 1990; Takada 1992 

Taiwan 

Chou 1981b, Takada 1992 

Trioxys nr pallidus Morocco Fulmek 1956 

Trioxys rietscheli China, India Shi 1980; Raychaudhuri 1990 

Trioxys rubicola India Agarwala et al. 1981 

Trioxys sinensis Pakistan Mohyuddin et al. 1972 

Trioxys sp. 

Portugal 

Taiwan, USA 

ENCYRTIDAE 

Aphidencyrtus sp. Malaysia Yunus & Ho 1980 

PTEROMALIDAE 

Pachyneuron aphidis China Shi 1987 

DIPTERA 

CECIDOMYIIDAE 

Aphidoletes aphidimyza Chile, Europe, 

Nth America 

Endaphis maculans Trinidad 

USA 


Bšrner et al. 1957; Boelpaepe et al. 1992 

Fulmek 1956 

Harris 1973; Kocourek et al. 1993, 

Meadow et al. 1985; Prado 1991 

Kirkpatrick 1954 

Tang et al. 1994; Yokomi et al. 1994 



ACARINA 


TROMBIDIIDAE 

Allothrombium pulvinum China Dong et al. 1992 

Xu et al. 1993 

Zhang et al. 1993; Zhang & Chen 1993 

*Starù (1967a, 1970) selected these species (bold type) for possible introduction to areas where they do not occur. 

Also recorded from Aphis craccivora 


4.4 Aphis gossypii 59 

Under humid conditions, high aphid mortality may result from fungal 

infection. The two species commonly reported are Neozygites fresenii and 

Cephalosporium (= Verticillium) lecanii (Table 4.4.2), although about a 

dozen species may be involved (Hagen and van den Bosch 1968). Effective 

use of the above fungi has been made under glasshouse conditions and 

V. lecanii is available commercially for this purpose. However, in Florida 

this fungus has performed poorly on A. gossypii compared with against 

Myzus persicae (Osborne et al. 1994). 

Table 4.4.2 Fungi attacking Aphis gossypii and/or A. craccivora 


Arthrobotrys sp. USA OÕBrien et al. 1993 

Beauveria bassiana USSR Pavlyushin & Krasavina 1987 

Cephalosporium lecanii Japan 

Netherlands 

USA 

USSR 

Venezuela 

Masuda & Kikuchi 1992; 

Saito 1988 

Yokomi & Gottwald 1988; 

Sopp et al. 1990; Vehrs & 

Parrella 1991; Schelt 1993 

Cermeli 1989 

Pavlyushin & Krasavina 1987 

Entomophthora exitialis India Kranz et al. 1977 

Entomophthora sp. Chile Prado 1991 

Neozygites fresenii Australia 

Chad 

China 

Cuba 

Milner & Holdom 1986 

Silvie & Papierok 1991 

Zhang 1987 

Hernandez & Alvarez 1985 

USA Steinkraus et al. 1991, 1995, 

1996; OÕBrien et al. 1993; 

Steinkraus & Slaymaker 1994; 

Smith & Hardee 1996 

Paecilomyces fumosoroseus USSR Pavlyushin & Krasavina 1987


Attempts at biological control 

There have been many intentional and unintentional transfers of aphid 

parasitoids, which have influenced the populations of A. craccivora and 

A. gossypii in different regions of the world. However, the majority of 

intentional transfers have been aimed at other target aphid pests. Some 

deliberate attempts against these two species have been unsuccessful (Table 

4.4.3). Overall, however, there is little doubt that, where parasitoids have 

become established, the situation is better, sometimes significantly better, 

than if they were not present, even if the level of control may not be as 

effective as is desirable. There is little doubt that, in many regions, an 

improved situation is likely to occur if additional parasitoid species are 

established. 

AUSTRALIA 

Aphidius colemani is capable of producing rapid decreases in populations of 

A. gossypii on cotton and, of carrying this to extinction in association with 

high densities of Harmonia octomaculata (= Coccinella arcuata) and 

Coccinella transversalis (= C. repanda) (Room and Wardhaugh 1977). 

Three species of parasitoid were imported in the hope that, as polyphagous 

species, they would contribute to the biological control of several species of 

pest aphid. The principal target for two of the species was A. craccivora, 

which is very sporadic in occurrence in Australia. It was hoped that, in the 

unpredictable absence of A. craccivora, the parasitoids would continue to 

breed and survive in other hosts. 

Lysiphlebus testaceipes (from Aphis nerii in California) and L. fabarum 

(from Greece and Turkey) were imported as biological control agents of 

A. craccivora on legumes, mass reared, and released in 1982 and 1983 in 

New South Wales and Victoria. Both parasitoids readily parasitised 

A. craccivora, A. gossypii and some other aphid species in the laboratory. 

The releases coincided with a prolonged drought during which there were no 

A. craccivora available on legume crops. Releases were, therefore, made on 

A. gossypii infesting Hibiscus bushes. No parasites were recovered the 

following year from either A. craccivora or A. gossypii, although 

L. testaceipes became established in Aphis nerii on oleander (Nerium) 

(Hughes 1989). L. fabarum was not recovered. It was concluded that the 

parasitoids must have been unsuitable biotypes (Carver 1984, 1989).

Table 4.4.3 Releases for the biological control (inter alia) of Aphis craccivora and/or A. gossypii 

Parasitoid From To Year Result Reference 

Aphelinus varipes South Carolina California + Wharton 1983 

Aphelinus abdominalis India U.K. + Hussey & Bravenboer 1971 

Aphidius colemani S. Brazil France 1982 + Rabasse 1986 

Australia Tonga 1990 + Carver et al. 1993; 

Wellings et al. 1994 

Lysiphlebus fabarum France, Italy, Greece, Australia 1982, 

- Carver 1984, 1989 

Turkey 

1983 

Lysiphlebus testaceipes USA 

China 

1983 

+ Zheng & Tang 1989 

USA 

Hawaii 1923 

+ Beardsley 1961 

USA 

India 

1966 

? Ramaseshiah et al. 1969; 

Sankaran 1974 

USA 

Australia 1982 

+ Carver 1984; Hughes 1989 

Cuba 

France, Italy 1973 

+ Rabasse 1986; Starù et al. 1988a,b 

Czechoslovakia Tonga 

- Stechmann & Všlkl 1988; Všlkl et al. 

1990 

Hawaii 

Pakistan 1972 

? Anwar 1974; 

Mohyuddin & Anwar 1972, 1973, 

Mohyuddin et al. 1971 

Mexico 

USSR 

1989 

+ Shiiko et al. 1991 

USA 

China 

1983 

+ Zheng & Tang 1989 

Trioxys indicus India Australia 1986 - Carver 1989; Sandow 1986 



CHINA 

Praon volucre was imported from the Mediterranean area for the 

biological control of Hyperomyzus lactucae, a vector of lettuce necrotic 

yellows. It was mass reared and released in 1981 and 1982, mainly in New 

South Wales and Victoria. It has not been recovered from mainland 

Australia, but is reported to be present in Tasmania. In laboratory trials it 

successfully parasitised a number of pest aphids that occur in Australia, 

including A. craccivora. 

In 1986 Trioxys indicus was introduced from India and released in 

Western Australia, Victoria and New South Wales against A. craccivora 

(Carver 1989), but establishment did not occur. 

Although he gives no further details Mohammad (1979) states that 

parasitisation of A. craccivora in Adelaide soon after colonisation 

frequently prevented the establishment of a colony. 

Zhang (1992) recorded for A. gossypii on cotton in China more than 48 

species of natural enemy (belonging to 19 families in 9 orders). Coccinellids, 

spiders and lacewings were the most important predators of this and other 

pests in cotton fields. Nan et al. (1987) record attack on A. gossypii and other 

cotton pests by 5 species of pentatomid, 9 species of coccinellid, 4 species of 

lacewing and 36 species of spider. The dominant predators studied by Wu 

(1986) were found to vary according to the season and included the 

coccinellids Coccinella septempunctata, Hippodamia variegata, Propylea 

japonica, Harmonia axyridis, the lacewings Chrysopa (= Chrysoperla) 

sinica, Chrysopa formosa, C. pallens (= C. septempunctata), C. intima and 

the spiders Erigonidium graminicolum, Misumenops tricuspidatus and 

Xysticus croceus. Zhang (1985) carried out laboratory tests on the daily 

consumption of A. gossypii by Scymnus hoffmanni, Chrysopa sinica and the 

spider Erigonidium graminicolum. Ma and Liu (1985) reported on the 

effectiveness and seasonal fluctuations of Propylea japonica and Yang 

(1985b) on its laboratory rearing. Ding and Chen (1986) examined the 

predation pattern of Chrysopa sinica. Propylea japonica, Scymnus 

hoffmanni and spiders (especially Theridion octomaculatum and 

Erigonidium graminicolum) were major enemies of A. gossypii on cotton in 

Hunan Province. The spiders (2.6 to 26 per 100 plants ) were present from 

late June to late August and were relatively unaffected by the weather. 

Coccinellid populations fluctuated somewhat with the season. Reproduction 

of A. gossypii was inhibited and its damage reduced when the ratio of total 

natural enemies to aphids was 1:50 or the ratio of coccinellids was 1:140. 

Since 1978, cotton fields over large areas have not been treated with 

insecticides before August, in order to safeguard the natural enemies which 

now hold the aphids in check (Mao and Xia 1983). Zhao et al. (1989) report


that the lycosid spider Pardosa astrigata is an important predator in cotton 

fields and Dong (1988) that the coccinellid Harmonia axyridis was an 

effective natural enemy when present in adequate numbers. Other predators 

include the anthocorid bug Orius minutus (Miao & Sun 1987), the 

coccinellids Scymnus hoffmanni (Zhao and Holling 1986), Propylea 

japonica and Harmonia axyridis (Zou et al. 1986; Lei et al. 1987 ). 

The aphidiid Aphidius picipes (= A. avenae) parasitised more than 80% 

of A. gossypii individuals on Chinese cabbage growing near cotton fields (Li 

& Wen 1988). Laboratory studies showed that Trioxys communis was more 

effective than Aphelinus mali in suppressing A. gossypii populations (Shi 

1985). Of the 5 species of parasitoid recorded by Xi and Zhu (1984) on 

A. gossypii on cotton in Jiangsu Province, Lysiphlebia japonica and Trioxys 

indicus were dominant and each accounts for about 45% of all parasitoids. In 

the laboratory, female L. japonica laid an average of 120 eggs and produced 

a parasitisation rate of up to 14%. In the field it overwinters as larvae inside 

A. gossypii, A. craccivora or Myzus persicae. In an earlier study Tian et al. 

(1981) recorded the same two parasitoids on both A. craccivora and 

A. gossypii with a combined parasitisation rate of 13%. This did not provide 

effective control and it was pointed out there was heavy attack by 

hyperparasitoids, such as Aphidencyrtus sp. (Encyrtidae). 

Larvae of the mite Allothrombium pulvinum have been observed 

attacking A. gossypii in cotton fields (Zhang and Chen 1993; Zhang et al. 

1993). 

COLOMBIA 

Aphelinus sp. caused 2.2% and Lysiphlebus sp. 0.3% parasitisation of 

A. gossypii on cotton in the field (Ramirez and Zuluaga 1995). 

CUBA 

Aphis craccivora is a common pest of vegetables and many other crops and 

also occurs on wayside trees, such as Gliricidia. In beans and other annual 

crops it occurs for a short period only, whereas on wayside trees it is present 

more or less continuously. The native parasitoid Lysiphlebus testaceipes 

parasitises the aphid heavily on Gliricidia, but poorly or not at all on young 

beans. This appears to be primarily a matter of the relative rates of dispersal 

of the aphid and its parasitoid (Starù 1970). 

EAST ASIA 

A list of parasitoids recorded from A. gossypii in East Asia is shown in table 

4.4.1 in which most entries from this region are based on Takada (1992). The 

most comprehensive information within this region is available from 

Taiwan (Starù and Schlinger 1967; Tao and Chiu 1971; Chou 1984), Japan 

(Takada 1968; Takada and Yamauchi 1979; Takada unpublished) and India 

(Raychaudhuri 1990). For Southeast Asia, there are two records from


FRANCE 

INDIA 

Vietnam and one from Malaysia. In so far as one can argue from such scanty 

data, the principal parasitoids of A. gossypii in the Far East (Trioxys 

communis, Lysiphlebia japonica and the particular species involved of 

Aphelinus) do not occur in India (Takada 1992). According to Takada, the 

principal species attacking A. gossypii in India is Trioxys indicus, which is 

recorded from Taiwan, but apparently not from Japan. Another species 

attacking A. gossypii in India is Lipolexis scutellaris, which also occurs in 

Vietnam, Malaysia and Hong Kong, but is not recorded elsewhere in the Far 

East (Takada 1992). Two widely distributed and effective parasitoids that 

attack A. gossypii in other parts of the world (Aphidius colemani and 

Lysiphlebus testaceipes) do not yet seem to be present in Southeast Asia, 

although A. colemani is recorded from the field in Pakistan (Starù 1975). 

Takada (1992) comments on the habitat specialisations in Japan of 

parasitoids of A. gossypii, which occurs in both open and lightly wooded 

habitats: Trioxys communis, and Aphelinus species prefer the open habitat, 

whereas Lysiphlebia japonica, Ephedrus nacheri, E. persicae and 

E. plagiator prefer the lightly-wooded habitat. Thus, the parasitoid complex 

on A. gossypii on cucumber, egg plant or taro is quite different from that on 

Hibiscus or Rhamnus in a garden. 

Starù et al. (1973) reviewed the parasitoids of aphids in France. A South 

American strain of Aphidius colemani, which is adapted to warm subtropical 

conditions and is highly polyphagous, was introduced from southern Brazil 

and released against Toxoptera aurantii in France near Antibes in 1982. It is 

reported to be established (Rabasse 1986; Tardieux and Rabasse 1986). In 

1973Ð74 Lysiphlebus testaceipes was introduced into France and released 

near Antibes and in Corsica. It was recovered soon after, and later in Italy. It 

was also sent to eastern Spain where it established and spread to become the 

predominant parasitoid in the regions where it occurs. It attacks A. gossypii 

on citrus and a number of other aphids on other host plants (Starù et al. 

1988a). 

Including the widespread Trioxys indicus, 14 parasitoids were recorded from 

A. gossypii (a preferred host) and 8 from A. craccivora (Agarwala 1983). 

Ephedrus persicae is reported to be confined in India to A. craccivora and 3 

parasitoids, Praon absinthii, Trioxys basicurvus and T. rubicola confined to 

the A. gossypii complex (Agarwala et al. 1981). 

The impact of the widespread parasitoid Trioxys indicus on 

A. craccivora feeding on pigeon pea was studied in the laboratory and the 

field. The parasitoid had a high searching ability and exhibited a density 

dependent relationship with its host. A single female oviposited in 100 to 

150 aphids in 3 to 5 days after emergence and the life cycle occupied 15 to 20

IRAQ 

ISRAEL 


days. Early in the season 9.4% of the aphids were parasitised, rising to a peak 

of 64.6% two months later and resulting in suppression of the aphid 

population. Trioxys indicus has a relatively narrow host range, which 

includes A. gossypii, A. craccivora and the oleander aphid A. nerii. Up to 

2.4% of the parasitoids were hyperparasitised by a cynipoid wasp. Singh and 

Sinha (1983) concluded from these studies that T. indicus had most of the 

necessary attributes of a potentially effective biological control agent for 

A. craccivora and A. gossypii. 

Ramaseshiah and Dharmadhikari (1969) found Aphelinus abdominalis 

(= A. flavipes) to be one of the important parasitoids of A. gossypii in India; 

furthermore, that Aphelinus sp. nr abdominalis had A. craccivora as a 

preferred host, but also attacked A. gossypii. 

Many generalist predators attack A. gossypii and other aphids in India, 

and Agarwala and Saha (1986) record 8 species attacking it there on cotton. 

Seven species of predatory coccinellid, 2 syrphids and a chrysopid were 

recorded preying on A. gossypii on potatoes. Coccinella septempunctata and 

Cheilomenes (= Menochilus) sexmaculata were the most important (Raj 

1989). A. gossypii was reported to be controlled on sunflower by 

coccinellids (Goel and Kumar 1990). The aphidophaghous coccinellid 

Micraspis discolor showed a preference for A. craccivora (Agarwala et al. 

1988). 

The biology of the predatory Leucopis species (Diptera) attacking both 

A. craccivora and A. gossypii on chrysanthemums is described by Kumar et 

al. (1988). Larvae of Chrysopa orestes had a substantial effect on A. gossypii 

on eggplant (Bhagat and Masoodi 1986). 

Raychaudhuri et al. (1979) record three predatory spiders: Cyclosa 

insulana (Araneidae) attacking A. craccivora and A. gossypii; Theridion sp. 

(Theridiidae) attacking A. craccivora; and Uloborus sp. (Uloboridae) 

attacking A. gossypii. 

A. gossypii is parasitised on melons, cotton and okra by Lysiphlebus 

fabarum which, in turn, is attacked by the hyperparasitoids Pachyneuron 

aphidis (Pteromalidae), Dendrocerus (= Lygocerus) sp. (Megaspilidae), 

Aphidencyrtus sp. (Encyrtidae) and Alloxysta (= Charips) sp. (Charipidae) 

(Al-Azawi 1966). 

A. gossypii occasionally infests citrus in the vicinity of cotton fields. Based 

on small samples, it was found to be attacked by only one parasitoid, Trioxys 

angelicae (Rosen 1967a,b).


ITALY 

A. gossypii in particular, but also A. craccivora, are two of the 10 aphid 

species attacking citrus. Eleven species of aphidiine parasitoids provide a 

considerable measure of biological control (Tremblay 1980). Lysiphlebus 

testaceipes (particularly) and L. fabarum are the commonest species and 

together may attain a parasitisation rate of 90% to 100%. 

The most important predators are coccinellids, of which Scymnus spp. 

are common amongst colonies of A. gossypii. Together with Coccinella 

septempunctata and other natural enemies, they may quickly suppress a 

cotton aphid population. Chrysopid, syrphid and cecidomyiid larvae are less 

effective although, in the absence of coccinellid larvae, syrphid larvae may 

be important. A. gossypii is under biological control on citrus in orchards 

where pest management procedures are adopted (Barbagallo and Patti 1983; 

Starù 1964). Recent papers on natural enemies of A. gossypii in Italy have 

been published by Ferrari and Burgio (1994) and Ferrari & Nicoli (1994). 

JAPAN 

Amongst numerous predators reported on A. gossypii by many authors are 

Coccinella septempunctata (Nozato and Abe 1988) and Scymnus hoffmanni 

(Kawauchi 1987). 

KOREA 

The consumption of A. gossypii by larvae of the coccinellid Harmonia 

axyridis was studied by Choi and Kim (1985). 

Eight species of parasitoid (and 6 of hyperparasitoid) of A. craccivora 

were reported by Chang and Youn (1983). The more important species (and 

rates of parasitisation) were Lysiphlebus fabarum (31.6%), Lipolexis 

scutellaris (18.8%), Lysiphlebia japonica (16.7%) and Adialytus salicaphis 

(11.4%). Of 509 field collected, mummified aphids 44.8% produced 

parasitoids and 43.8% hyperparasitoids giving an overall parasitisation rate 

of 88.6%. 

MALAYSIA 

The only record of parasitoids of A. craccivora or A. gossypii appears to be 

that of Ng and Starù (1986). Lipolexis scutellaris, an oriental species with a 

wide distribution in India and extending to Vietnam, southern China and 

Taiwan, was recorded from both aphid hosts. Trioxys communis, also an 

oriental species, was rarely found, but only on another aphid species (Aphis 

spiraecola). At least 3 unidentified species of aphelinids were bred and in 

large numbers, but no other information on these is provided. 

NETHERLANDS 

A valuable review of the biological control of A. gossypii in glasshouses, 

with special reference to the situation in the Netherlands, is provided by van 

Steenis (1992). van Steenis (1995) evaluated 4 aphidiine parasitoids for 

biological control of A. gossypii on glasshouse cucumbers. Aphidius 

colemani performed the best (72 to 80% parasitisation), followed by


Lysiphlebus testaceipes (26%), Ephedrus cerasicola (23%) and Aphidius 

matricariae (less than 5%). The general principles of selection and 

establishment of species are relevant elsewhere, although only the first of the 

three species selected (Lysiphlebus testaceipes, Aphidius matricariae, 

Ephedrus cerasicola) would appear to be particularly relevant to Southeast 

Asia or the Pacific. 

PAKISTAN 

In investigations from 1967 to 1970, 6 parasitoid species and 22 predator 

species were recorded attacking Aphis craccivora on a range of leguminous 

crops and weeds in 5 climatologically different zones of Pakistan (Hamid et 

al. 1977). Alate A. craccivora were found on soybean (Glycine max), but 

colonies of apterous aphids never developed, possibly due to the presence of 

abundant plant hairs. 

Aphelinus abdominalis (= A. basalis), which was widespread and active 

throughout the year, parasitised from 0.6% to 17.6% of A. craccivora, the 

level depending upon host plant, location (hills, foothills or plains) and 

season. Perhaps due to its small size, A. abdominalis was the only parasite 

that attacked A. craccivora under the covering of hairs on Phaseolus aureus. 

The period from oviposition in the aphid to mummy formation was 3 to 4 

days and adult wasps then emerged in 5 days. Trioxys ?sinensis parasitised 

up to 21.3% of aphids. Development from oviposition in the aphid to 

mummy formation took 14 to 16 days and adults emerged 4 to 7 days later. 

Lysiphlebus fabarum parasitised between 1.3% and 72.9% of available 

hosts, Ephedrus nr cerasicola 1.3% to 1.4% and Aphidius absinthii 11%. A 

low level of attack on most of the parasitoids was recorded by the 

hyperparasitoids Alloxysta sp. and Pachyneuron sp. 

Ants (Monomorium indicum and Pheidole sp.) were associated with over 

90% of the aphid colonies. The size of A. craccivora on Vicia faba nursed by 

ants was far greater than those not attended and aphid mortality was higher 

when ants were absent. The most abundant of 22 predator species (Table 

4.4.2) were Cheilomenes sexmaculata, Coccinella septempunctata and 

Syrphus spp. 

In the same environment Trioxys sinensis and Lysiphlebus fabarum 

parasitised Aphis gossypii on cucumbers and on Hibiscus esculentus (Hamid 

et al. 1977). 

PHILIPPINES 

A. craccivora is commonly attacked by Lipolexis scutellaris; and A. gossypii 

by this species and also Trioxys communis. Both parasitoids also attack other 

species of aphids, but not the banana aphid Pentalonia nigronervosa (V.J. 

Calilung pers. comm. 1995).


RƒUNION 

A. craccivora is parasitised by Aphidius colemani on Gliricidia maculata 

and by Aphelinus sp. on Vigna unguiculata. In turn Aphelinus sp. is 

parasitised by Syrphophagus africanus and Pachyneuron vitodurense. 

A. craccivora is also attacked by the coccinellid predators Scymnus 

constrictus and Platynaspis capicola (Quilici et al. 1988). 

SHANGHAI 

The main parasitoids of A. gossypii on cotton were Trioxys communis, 

T. rietscheli and Lipolexis gracilis. Next in importance was Aphidius 

gifuensis, and there was occasional attack by Aphelinus abdominalis and 

A. mali. The highest total parasitisation recorded was about 27%. Parasitoids 

constituted 22.7% of the emergences from aphid mummies and the 

hyperparasitoids Syrphophagus aphidivora 45.2%, Alloxysta sp. 15.1%, 

Pachyneuron aphidis 14.7% and Dendrocerus 2.3%. The number of 

parasitoids only exceeded that of hyperparasitoids during the first half of 

August (Shi 1980, 1987). 

TONGA 

Aphidius colemani and Lysiphlebus testaceipes were introduced from 

cultures in Czechoslovakia for the biological control of the banana aphid 

Pentalonia nigronervosa (Stechmann and Všlkl 1988, 1990; Všlkl et al. 

1990), but there is no indication that they became established. A. colemani 

from a culture originating from a garden in Canberra was introduced again in 

1990 and recovered in 1992 from Aphis gossypii on taro, but not from the 

banana aphid. The further introduction of Aphidiidae, which are obligate 

parasitoids of aphids, was recommended since they could assist in the 

control of pest aphids and not pose a threat to non-target insects (Carver et al. 

1993). Although 15 aphid species are recorded in Tonga including 

A. craccivora, by 1993 no aphids other than A. gossypii had been recorded 

as hosts of A. colemani, although recent monitoring has not been possible. 

Two other primary parasitoids were recorded, Aphelinus gossypii (from 

Aphis gossypii) and Lipolexis scutellaris, from a single female, free on a 

banana sucker (Carver et al. 1993; Wellings et al. 1994). 

Three common and widespread aphid predators were recorded (the 

syrphid Ischiodon scutellaris, the coccinellid Harmonia octomaculata and 

the hemerobiid Micromus timidus) in addition to 11 tramp species of ants 

(Carver et al. 1993).

USA 

USSR 


Lysiphlebus testaceipes parasitised 74.5% of Aphis gossypii on strawberries 

and Aphelinus semiflavus a smaller number. Seven hyperparasitoids were 

reared (Oatman et al. 1983b). L. testaceipes was considered by Schlinger 

and Hall (1960) to be the most effective aphid parasitoid in southern 

California and to give excellent control there of Aphis gossypii. At least 8 

hyperparasitoid species were also reared (Schlinger and Hall 1960). 

Entomopathogen infection was the primary cause of a reduction in 

A. gossypii population that occurred during the week after peak aphid 

abundance on cotton in Mississippi and continued pathogen activity, 

combined with predation, maintained aphids at a low density for the 

remainder of the season. Early in the season parasitisation and predation 

may have reduced aphid population growth (Weathersbee and Hardee 1993, 

1994). 

In untreated cotton plots small predators (spiders and Geocoris spp.: 

Hemiptera, Lygaeidae) had the greatest impact on A. gossypii populations 

and the parasitoid Lysiphlebus testaceipes was never abundant. Fungi killed 

many aphids and constituted the most important natural enemy factor in 

insecticide treated plots (Kerns and Gaylor 1993). Fungi attacking 

A. gossypii in USA include Neozygites fresenii (Steinkraus et al. 1992, 

1993a,b,c; Sanchez-Pena 1993, Smith and Hardee 1993) and 

Cephalosporium (= Verticillium) lecanii (Sopp et al. 1990, Yokomi and 

Gottwald 1988). The coccinellid predators Hippodamia convergens and 

Scymnus louisianae, the chrysopid Chrysoperla carnea, and Syrphus sp. 

were effective in reducing populations of A. gossypii in Texas (Vinson and 

Scarborough 1989). In Alabama the hemerobiid Micromus posticus is an 

important predator (Miller and Cave 1987). 

In southern USSR coccinellid beetles are important predators of aphids. 

Adult Coccinella undecimpunctata and larvae of Hippodamia variegata are 

the most voracious and prefer A. gossypii, whereas Coccinella 

septempunctata prefers A. craccivora (Belikova and Kosaev 1985). 

There are many papers dealing with the control of A. gossypii and 

associated pests in glasshouses. The lacewing Chrysoperla carnea was 

effective only when released at a predator: aphid ratio of 1:20, whereas 

Chrysopa sinica was effective at 1:50 (Shuvakhina 1983). Other predators 

utilised include Chrysopa perla (Ushchekov 1989) and the cecidomyiid 

Aphidoletes aphidimyza (Begunov and Storozhkov 1986). Under conditions 

of high humidity, high aphid mortality was caused by the fungi 

Cephalosporium lecanii, Beauveria bassiana and Paecilomyces 

fumosoroseus (Pavlyushin and Krasavina 1987).


VIETNAM 

A. gossypii was one of four aphids surveyed for parasitoids by Starù and 

Zelenù (1983), but the number of aphidiid species found (2) was surprisingly 

low. Lipolexis scutellaris was commoner than Lysiphlebia mirzai. The 

former parasitises a number of other Aphis species, including Aphis 

spiraecola (= A. citricola), A. craccivora and A. nerii. Some unidentified 

aphelinid parasitoids were also present. 

Starù and Zelenù (1983) suggest that Lipolexis scutellaris may be a 

valuable species for transfer elsewhere and also that Vietnam would benefit 

from the introduction of additional parasitoid species. 

The major parasitoid species 

General features of the Aphidiidae 

Different populations of many Aphidiidae have differing biological 

properties and are often known as biotypes, i.e. contrasting groups, each 

consisting of individuals of the same species. Biotypes are recognised by 

biological function rather than morphology and consist of those individuals 

that behave similarly as far as our immediate interests are concerned 

(Mackauer and Way 1976). 

The members of this family attack aphids exclusively and are probably 

the most commonly observed cause of aphid mortality in the field. In 

Europe, many, if not all, aphid colonies come to include some mummified 

individuals (i.e. dead aphids containing a fully-grown parasitoid larva or 

pupa) (Starù 1970). Aphidiidae may hibernate as prepupae within host 

mummies. All are solitary endoparasitoids. All aphid stages are attacked 

except the eggs, but alatae are least often attacked. The parasitoid egg is 

usually inserted anywhere in the aphid abdomen. The preferred aphid larval 

instar varies with the parasitoid species, but younger instars are usually 

chosen. If adult aphids are parasitised the parasitoid larva may not complete 

its development before the insect dies, so that the parasitoid perishes also. 

Oviposition into an aphid does not ensure the successful development of a 

parasitoid, since the host may be unsuitable or it may already be parasitised: 

defence and immune responses are common. However parasitised hosts are 

usually distinguished by the parasitoid and receive no further eggs. 

If an aphid is parasitised during the last larval instar, a mummy is formed 

after it moults to the adult. The first and second instars of aphidiid 

parasitoids generally feed on haemolymph, but the last instar attacks the 

alimentary canal and other organs, ultimately killing its host. The parasitoid 

larva then spins a cocoon and pupates inside the empty aphid skin. The adult 

emerges through a small circular hole usually cut dorsally or apically near


the posterior of the mummy. Aphidiid mummies are round and usually 

straw-coloured to brown and in some genera (e.g. Ephedrus) always black 

and parchment-like. Aphelinid larvae do not spin cocoons, and their 

mummies are usually slender and black (Takada 1992). 

The chain of events that determines host specificity includes, in 

sequence, host habitat finding, host finding, host acceptance by the 

parasitoid and host suitability. 

The last larval instar of some parasitoids provokes their aphid hosts to 

move away from the plant on which they were feeding. With Lysiphlebus 

fabarum, Ephedrus plagiator and Trioxys angelicae this migration of premummies 

is connected with diapause (under conditions of a short day) or 

with aestivation (under conditions of a long day). Parasitoids usually emerge 

without delay from aphids which become mummies where they have been 

feeding (Behrendt 1968). 

Starù (1970) provides additional details of many of the species of 

Aphidiidae that follow below and HŒgvar and Hofsvang (1991) a 

comprehensive review of their biology, host selection and use in biological 

control. 

Aphelinus abdominalis (= A. flavipes) Hym.: Aphelinidae 

This species is widespread in Europe and is recorded also from USSR, India, 

Australia and Israel. It is extensively distributed in India as one of the 

important parasitoids of Myzus persicae and A. gossypii. It becomes active 

in late May and is abundant during June and July. The incubation time for 

eggs is 2 days and adults emerge after 13 days in September (Ramaseshiah 

and Dharmadhikari 1969). 

When an Indian strain was liberated in a U.K. glasshouse at 23¡C, one 

week after artificially infesting plants with A. gossypii, it was unable to 

overtake the pest population because the rate of increase of the pest was 

scarcely affected by the parasitoid. Only when aphid overcrowding occurred 

and rate of increase was self-limited, did the parasiteÕs rate of increase (6 ´ per 

week) exceed that of the aphid. Reducing the glasshouse temperature to 19¡C 

slowed the rate of aphid increase and permitted the parasitoids to contain the 

pest before severe leaf-distortion occurred. On the other hand, when 

parasitoids were present at the time that A. gossypii was introduced, aphid 

reproduction was suppressed and effective control resulted (Hussey and 

Bravenboer 1971). 

Aphelinus gossypii Hym.: Aphelinidae 

This species was described from Hawaii and is recorded from Australia, 

New Zealand, India, Japan and also from Tonga, where it was reared in 

abundance from Aphis gossypii on taro (Colocasia esculenta). It is probably


present elsewhere under other names (Carver et al. 1993). Aphelinus 

gossypii lays in some host eggs encountered, but also kills many others by 

probing and then feeding on exuding fluids (Takada and Tokumaku 1996). 

Parasitisation is reduced when the host is protected by the presence of ants 

(Stechman et al. 1996). Aphelinus gossypii was parasitised in Tonga to the 

extent of 30% to 60% by the cynipoid hyperparasitoid Alloxysta darci. This 

parasitises species of Aphelinus, but not of Aphidiidae so it is most unlikely 

to parasitise Aphidius colemani, a recently established parasitoid attacking 

Aphis gossypii there. Alloxysta darci was earlier incorrectly identified as 

Alloxysta brevis (Carver 1992; Carver et al. 1993). Aphelinus gossypii is an 

effective parasitoid at low A. gossypii density and hence an important 

candidate for consideration for introduction (P. Wellings, pers. comm.). 

Aphelinus mali Hym.: Aphelinidae 

This well known North American parasitoid of above-ground stages of the 

woolly apple aphid Eriosoma lanigerum has been introduced intentionally 

or inadvertently into almost every country where its host has established 

itself as a pest. It is a very effective parasitoid in moderately warm climates 

(Rosen 1967b) and has occasionally been reported from other hosts, 

including Aphis gossypii, although in such instances it has probably been 

confused with the very similar Aphelinus gossypii (M. Carver pers. comm.). 

The life cycle occupies almost 20 days in summer (egg 3, larva 10 to 12 and 

pupa 6 to 7 days respectively). Parasitised aphids have a strong tendency to 

seek sheltered places before death (Clausen 1978). 

Aphelinus semiflavus Hym.: Aphelinidae 

This species has a wide host range and occurs in USA, Hawaii, India and 

Europe. It overwinters as a pupa in its host. Single eggs are laid, generally in 

the dorsal surface of the host abdomen. Young aphids are preferred hosts, but 

even adults are parasitised, in which case fewer young are produced by the 

aphid before it is killed. Over 600 eggs may be produced by a female which 

often feeds on haemolymph exuding from oviposition punctures. Males are 

rare and females can produce offspring parthenogenetically. Developmental 

periods are: egg 3 days, larva 6 to 11 days and pupa 7 to 8 days. Although 

Myzus persicae is preferred, A. semiflavus also parasitises Aphis gossypii as 

one of 15 or so other hosts. In USA it was attacked by 3 hyperparasitoids, 

Asaphes lucens (= A. americana) (Pteromalidae), Alloxysta sp. (Charipidae) 

and Syrphophagus aphidivora (= Aphidencyrtus aphidiphagus) (Encyrtidae) 

(Hartley 1922; Ramaseshiah and Dharmadhikari 1969; Schlinger and Hall 

1959).


Aphidius colemani Hym.: Aphidiidae (= A. platensis, 

= A. transcaspicus) 

Starù (1975) postulated that this species originated in India or nearby 

(possibly the Eastern Mediterranean). It is now widely distributed in 

Mediterranean Europe, Asia Minor, Central Asia, India, Pakistan, Africa, 

South America, Australia, New Zealand and New Caledonia (Starù 1972). In 

addition there have been intentional introductions to California, U.K., 

Czechoslovakia, Kenya (Starù1975) and Tonga (Carver et al. 1993). It is 

rather strange, if it originated in India, that it appears to be absent from 

Japan, China and possibly some of Southeast Asia (Starù 1975; Takada 

1992). A. colemani is restricted to the family Aphididae. Hosts consist of at 

least 9 species of Aphis, including A. craccivora and A. gossypii and at least 

30 species in other genera (Elliott et al. 1994; Starù 1975). 

In the field in Australia A. colemani is known to parasitise many species 

in the aphid tribes Aphidini and Myzini, but rarely species in the 

Macrosiphini and even more rarely species in other subfamilies (Carver et 

al. 1993). 

There are significant differences between countries both in the range of 

hosts attacked by A. colemani and the preference for particular host species 

(e.g. Messing and Rabasse 1995). This indicates that the species that is 

known as A. colemani is a complex of closely-related species or biotypes. 

For example, A. colemani parasitises Melanaphis donacis and Hyalopterus 

pruni in Mediterranean Italy and France, but none of the many other aphids 

present; in Central Asia only the latter aphid is attacked and in Iraq both 

aphids are attacked, in addition to Aphis zizyphi and A. punicae. An Italian 

population from Hyalopterus pruni was successfully reared on both Aphis 

craccivora and A. fabae in the laboratory. Furthermore, a French population 

from Melanaphis donacis was readily reared in the laboratory on Aphis 

craccivora, A. fabae and Myzus persicae (Starù1975). As another example, a 

strain (from Brazil) of Aphidius colemani successfully parasitised the 

oleander aphid Aphis nerii in France whereas another strain (from France) 

failed to do so (Tardieux and Rabasse 1986, 1988). In Mediterranean regions 

A. colemani parasitised A. gossypii successfully at 20¡C, but at temperatures 

above 27¡ it frequently failed to do so (Guenaoui 1991). The number of eggs 

laid per female A. colemani was 302 at 20¡C and 388 at 25¡C and 

development time to adult was 12.7 days and 10.0 days respectively. The 

intrinsic rate of increase of the parasitoid was similar to that of A. gossypii, 

suggesting that it is a promising parasitoid (van Steenis 1993). The optimum 

scheme for introducing A. colemani into glasshouses for control of 

A. gossypii on cucumbers has been investigated by van Steenis et al. (1996). 

Chou (1984) recorded A. colemani from TaiwanÑthe first record of this 

species from east AsiaÑwith H. pruni as its only host.


The care needed in selecting a biotype appropriate to the target pest is 

also illustrated by the following example. A. colemani from Aphis nerii 

mummies on a garden plant (Tweedia coerulia) in Canberra was readily 

reared for some generations in an insectary on the banana aphid Pentalonia 

nigronervosa before release in Tonga for biological control of that species. It 

has not been recovered from the banana aphid, but is now well established on 

A. gossypii attacking cucurbits (Carver et al. 1993; Wellings et al. 1994). 

When P. nigronervosa colonies are small they are mainly located deep in the 

leaf sheaths and they only extend into more exposed areas as they increase in 

size. Stadler and Všlkl (1991) found that Lysiphlebus testaceipes searched 

mainly in exposed areas for hosts, whereas A. colemani searched both 

exposed and concealed areas. This suggests that A. colemani would be the 

more appropriate of the parasitoids for hosts in concealed situations and, 

interestingly, it has been reported from P. nigronervosa in the field in 

northern New South Wales (M. Carver pers. comm.), where it is rare and was 

not encountered in recent searches (P.W. Wellings pers. comm.). 

A. colemani (often under one of its synonyms) has been introduced to 

several countries for the biological control of a range of aphid species (Starù 

1975). 

Aphidius gifuensis Hym.: Aphidiidae 

This species is native to the Oriental region. Details of its fecundity, 

oviposition period and longevity are provided by Fukui and Takada (1988). 

Aphidius matricariae Hym.: Aphidiidae (= A. phorodontis) 

This species is native to the temperate zones of the Palearctic region and has 

been recorded from more than 40 aphid species in Europe, North Africa, the 

Middle East, Israel, Mongolia and North and South America. It has a 

preference for the green peach aphid Myzus persicae in Israel (Rosen 

1967a,b) and California (Schlinger and Mackauer 1963). After contact with 

honeydew or an aphid host the time spent in searching that region for hosts 

increased (Masum 1994). 

Ephedrus persicae Hym.: Aphidiidae 

This is an almost cosmopolitan species, which is probably native to the 

Middle East or Central Asia, and now occurs in the Far East, Europe, South 

Africa, Madagascar, Australia and North America. It prefers leaf-curling 

aphid hosts, mainly belonging to the Myzinae and, less frequently, to the 

Aphidinae (Aphis spp.) (Mackauer 1963, 1965; Starù 1966). A review of the 

taxonomy and biology of Ephedrus persicae and E. plagiator is provided by 

GŠrdenfors (1986).


Ephedrus plagiator Hym.: Aphidiidae 

This aphid is native to the far eastern deciduous forests and steppes of the 

Palearctic region and is widely distributed in India. It has many hosts 

amongst species of Aphis and Myzus (Starù 1967a). 

Lipolexis gracilis Hym.: Aphidiidae 

This is a European or Far Eastern species with hosts in a number of aphid 

genera, including Aphis (Starù 1967a). 

Lipolexis scutellaris Hym.: Aphidiidae 

This is an oriental species (Raychaudhuri 1990) and is known from southern 

China, Japan and Taiwan and also from India, Pakistan and Tonga. It has a 

wide host range and an apparent preference for Aphis species (Carver et al. 

1993). 

Lysiphlebia japonica Hym.: Aphidiidae 

This is native to the Far East and is probably a well-adapted species for 

tropical climates. It typically occurs in forest or open woodland 

environments on many species in the genus Aphis in addition to those in a 

number of other related aphid genera (Starù 1967b). 

Lysiphlebia mirzai Hym.: Aphidiidae 

This species was described from India and is known also from Vietnam and 

China. 

Lysiphlebus fabarum Hym.: Aphidiidae 

(= Lysiphlebus confusus, L. ambiguus) 

This is a Palaearctic species (Europe, Asia Minor, Caucasus, Central Asia), 

which is now widespread and occurs also in Israel, a number of African 

countries and USA. It is the most abundant parasitoid of the black citrus 

aphid Toxoptera aurantii in Italy (Starù 1964) and Israel (Rosen 1967a,b). In 

some countries it is biparental but, in Israel, only females are known (Rosen 

1967a,b). It is recommended by Starù (1967b) as a species useful for 

biological control. L. fabarum is both biparental and parthenogenetic 

(Carver 1984) and 15 to 16 generations a year have been recorded in Italy 

(Tremblay 1964). It has a very extensive host range, with records from at 

least 144 species of aphids in 36 genera, 81 (56%) of these species belonging 

to the genus Aphis (Carver 1984). 

There is little doubt that L. fabarum refers to a complex of closely related 

sibling species or at least of host-specific biotypes. For example, in the 

laboratory L. fabarum bred from Aphis species readily parasitised other 

Aphis species, but not Brachycaudus sp.. However, in the field, colonies of 

Brachycaudus cardui heavily parasitised by L. fabarum shared the same 

host plants as unparasitised Aphis fabae (Mackauer 1962a). The influence of


temperature and humidity on the development of L. fabarum in A. gossypii 

and A. craccivora has been reported by Davletshina and Gomolitskia (1975) 

and methods for its mass production by Tregubenko and Popushoi (1987). 

Lysiphlebus testaceipes Hym.: Aphidiidae 

This parasitoid has a natural range extending from North America through 

Central America to the northern parts of South America. It is now known 

also from Hawaii, Australia, Europe and East Africa. It is the commonest 

native parasitoid of aphids in Mexico (Starù and Remaudi re 1982). It has a 

very wide host range, having been reported from at least 79 aphid species (32 

in the genus Aphis) in 32 genera (Carver 1984). 

Oviposition generally occurs in the abdomen of half grown and 

unparasitised hosts and, when hosts are scarce, more than one egg may be 

deposited (Sekhar 1957). Up to 254 eggs may be laid. It is heavily attacked 

by hyperparasitoids in its natural range (eg. 6 species when attacking Aphis 

gossypii (Schlinger and Hall 1960) or 7 species (Oatman et al. 1983b)). 

L. testaceipes was present in Australia (New South Wales and South 

Australia) prior to its introduction as a biological control agent and attacked 

an indigenous aphid Aphis acaenovinae (Starù and Carver 1979). 

There are many examples to demonstrate that L. testaceipes consists of 

biotypes. Thus, Californian L. testaceipes is unable to complete its 

development on Aphis spiraecola, whereas a Cuban strain does so 

successfully. The Californian strain did not attack Aphis nerii after 

introduction to Hawaii, although a Mexican strain subsequently introduced 

did so (Starù 1970). Then again, the biotype from A. craccivora on Robinia 

pseudacacia does not parasitise this same aphid on Phaseolus vulgaris; 

another biotype prefers A. gossypii on squash to this same aphid on hibiscus 

(Sekhar 1960; Tremblay and Barbagallo 1982). The effectiveness of 

Lysiphlebus testaceipes as a parasitoid on A. gossypii is thus significantly 

affected by the host plant on which the aphid is feeding (Steinberg et al. 

1993). 

L. testaceipes is reported to attack A. gossypii in Cuba (Starù 1981), 

Mexico (Starù and Remaudi re 1982) and, after introduction to Europe, in 

Spain, France and Italy. In Europe it now attacks more than 26 aphid species 

including A. craccivora and A. gossypii, often with high levels of 

parasitisation (Starù et al. 1988a,b,c). 

Lysiphlebus testaceipes was successfully introduced in 1923 from 

California to Hawaii for the biological control of aphids, including Aphis 

craccivora and A. gossypii. It soon spread widely throughout the islands, 

attacking these and other aphid species. In 1965, a further introduction from 

Mexico was made to control the oleander aphid Aphis nerii which had 

previously escaped attack. By 1927 several hyperparasitoids were recorded


as attacking L. testaceipes breeding in Rhopalosiphum maidis, an important 

virus vector on sugarcane (Timberlake 1927). According to Starù (1970), the 

introductions were partially to substantially effective, but importation of 

additional species was recommended. L. testaceipes from Cuba was 

introduced to Czechoslovakia for the biological control of aphids in 

greenhouses and also of some pest aphids in some subtropical areas (Starù 

1970). L. testaceipes was introduced in 1956 and 1960 from Hawaii to the 

Philippines, but no recoveries have been recorded (Baltazar 1963). 

L. testaceipes was introduced in 1973 from Cuba into France and Corsica 

(Italy) to reduce the numbers of citrus aphids (Starù et al. 1988b). It became 

well established, heavily parasitising Aphis gossypii and several other aphids 

(Rabasse 1986). 

Although there do not seem to be comparable data for A. gossypii, Hall 

and Ehler (1980) found that L. testaceipes averaged 79.5% parasitisation of 

Aphis nerii populations on oleander, with an average density of 12.4 aphids 

per shoot. When natural enemies were excluded, an average of 32.6 aphids 

were present per shoot, a clear indication that parasitisation was having a 

significant effect. A hyperparasitoid Pachyneuron sp. was active, but 

appeared to be generally unimportant in aphid population regulation. The 

average fecundity of L. testaceipes was found to be 128.2 eggs at 20¡C and 

180 eggs at 25¡C. Development from egg to female adult was completed in 

12.9 days at 20¡C and 9.5 days at 25¡C; and the life span of females was 2.7 

and 2.6 days at 20 and 25¡C respectively. At 20¡C the intrinsic rate of 

increase was slightly lower than that of Aphis gossypii, but was the same at 

25¡C. It was concluded that, at temperatures below 25¡C, L. testaceipes 

might not be able to overtake an established population of A. gossypii (van 

Steenis 1994). Earlier Schlinger and Hall (1960) concluded that 

L. testaceipes was capable of producing excellent control of both 

A. craccivora and A. gossypii. 

Praon volucre Hym.: Aphidiidae 

This is a palearctic species and is known from the Middle East, North Africa, 

India and Central Asia. It has an extensive and diverse host range, having 

been recorded from at least 90 aphid species in 35 genera. There is good 

evidence that P. volucre exists as a complex of host-specific biotypes or 

sibling species (Mackauer 1959, 1962a,b). Its biology has been studied by 

Beirne (1942). As in other Praon spp., pupation takes place under the empty 

mummy of the parasitised host. P. volucre females have been observed to 

use their front legs to hold the host aphid during oviposition (Beirne 1942). 

Although it has been recorded from A. craccivora in the field (Starù 1967a) 

and the laboratory (Carver 1984), it does not seem to have been reported 

from A. gossypii.


Trioxys angelicae Hym.: Aphidiidae 

This is widely distributed in Europe, Asia Minor and North Africa and has 

been reared from a wide range of hosts (Rosen 1967a,b). 

Trioxys communis Hym.: Aphidiidae 

This very important parasitoid of A. gossypii in China has been studied in a 

series of papers by Shi (1984, 1985, 1986). It develops in 8 days at 30¡C and 

in 16 days at 20¡C. When 4th instar A. gossypii are parasitised they 

mummify in the adult stage, having produced very few offspring. It is 

hyperparasitised by the pteromalid Pachyneuron aphidis. 

T. communis has also been recorded from A. gossypii in Japan, Korea, 

Taiwan and India and has been taken rarely in Malaysia from Aphis 

spiraecola (= A. citricola), but apparently not from other aphids (Ng and 

Starù 1986). 

Trioxys indicus Hym.: Aphidiidae 

A valuable review of the biology, ecology and control efficiency of this 

parasitoid is presented by Singh and Agarwala (1992). Its biology has been 

studied by Subba Rao and Sharma (1962) and later in a long series of papers 

from India (e.g. Singh et al. 1979; Sinha and Singh 1980a,b; Singh and Sinha 

1980a,b,c, 1982a,b; Pandey et al. 1982, 1984; Kumar et al. 1983; Singh and 

Pandey 1986; Singh and Srivastava 1988a,b, 1991, Singh and Agarwala 

1992). Ghosh and Agarwala (1982) provide host, host plant records and 

information on its distribution in India. 

Recorded hosts of T. indicus belong to 24 species of aphids in 14 genera, 

of which species of Aphis are best represented. The majority of host aphids 

are polyphagous, 6 are oligophagous and 1 monophagous. T. indicus prefers 

hosts on cultivated and wild shrubs to those on herbaceous and woody 

plants. A. craccivora is parasitised to the extent of 87% on pigeon pea (Singh 

and Tripathi 1987) and A. gossypii to the extent of 60% on both bottle gourd 

(Singh and Bhatt 1988) and eggplant (Subba Rao and Sharma 1962), and 

30% on cotton (Agarwala 1988). 

Based on extensive field observations, the three main hosts of T. indicus 

are A. craccivora, A. gossypii and A. nerii, each of which is parasitised by a 

range of other polyphagous parasitoids. 

The native range of T. indicus is largely the Indian subcontinent, 

although it has also been recorded from Taiwan (Starù and Schlinger 1967). 

It is most abundant in tropical and subtropical regions, where its numbers are 

comparatively low in summer and in rainy months and higher in the cooler 

months (Agarwala 1988). 

T. indicus females prefer to oviposit in second and third instar host 

nymphs. Probing without oviposition is common with first and second

Diptera 


instars, leading to high aphid mortality (up to 80% for first instars). Hosts 

that are already parasitised are generally avoided. Fecundity varies, but the 

figure of 143 offspring per female is quoted. The average time from 

oviposition to emergence is 10 days at 24 to 27¡C on A. gossypii (Subba Rao 

and Sharma 1962) and 16 to 18 days at 24 to 26¡C on A. craccivora. 

T. indicus lives less than 10 days in the laboratory (Pandey et al. 1982). 

Augmentation of T. indicus early in the season in pigeon pea fields in India 

was sufficient to control A. craccivora (Singh and Agarwala 1992). Extracts 

of A. craccivora sprayed on pigeon pea increased the fecundity of T. indicus 

and reduced the population doubling time (Singh and Srivastava 1991). 

A density-dependent relationship between T. indicus and its hosts has 

been reported by several authors (Subba Rao and Sharma 1962; Singh and 

Sinha 1980a; Saha and Agarwala 1986; Bhatt and Singh 1991a,b ). 

Eleven hyperparasitoids of T. indicus are known (Singh and Agarwala 

1992) and these should be rigorously excluded in any biological control 

transfers. 

Singh and Agarwala (1992) conclude that T. indicus is a very important 

parasitoid, especially of several species of Aphis in India, that it possesses 

most of the desirable attributes of a successful biological control agent and is 

therefore a promising natural enemy for introduction elsewhere against 

relevant pest aphids. It is also of value for inundative releases (Singh and 

Rao 1995). 

Endaphis maculans Dip.: Cecidomyiidae 

This aphid endoparasitoid attacked Toxoptera aurantii freely, but 

A. gossypii only lightly. It was seldom found in A. craccivora (Kirkpatrick 

1954). E. maculans lays its eggs on aphid-infested leaves and, upon 

hatching, the larva searches for aphids. When a host is encountered the larva 

penetrates the aphids dorsum and develops as an endoparasitoid, leaving as a 

mature larva via the aphidÕs anus. Average development time from egg to 

adult at 25¡ to 26¡C was 19.1 days (Tang et al. 1994). 

An aphid-specific predator 

Aphidoletes aphidimyza Dip.: Cecidomyiidae 

This is a common and widely distributed species throughout the northern 

hemisphere. It has been recorded from Japan, USSR, Czechoslovakia, 

Austria, Germany, Finland, France, Netherlands, U.K., Italy, Israel, Egypt, 

Sudan, Canada, USA and Hawaii. It is not recorded from Australia or New 

Zealand.


Comments 

Larvae of Aphidoletes reportedly feed exclusively as predators on aphids 

and are hence more host-specific than many of the other predators: 

A. aphidimyza is the best known of the cecidomyiid predators. Adults 

emerge during the day from pupae in the soil. They generally fly between 

sunset and sunrise. Orange-coloured eggs are laid singly or in clusters of up 

to 40, usually on plants near aphid colonies. Females live up to 14 days in the 

laboratory and lay about 100 eggs. These hatch after 3Ð4 days and first instar 

larvae immediately seek out and attack aphids. They usually attack by 

piercing a leg joint or some other joint. A toxin is perhaps injected, since the 

aphid is rapidly immobilised before its body fluids are extracted. The 

shrivelled bodies of some aphids remain attached to the plant by the stylet. 

Larval development involves 3 instars and takes 7 to 14 days (Harris 1973). 

On the other hand, Herpai (1991) reports 21 days from egg to adult (egg 3 

days, larva 8 days, pupa 10 days). 

Roberti (1946) gave a figure of 60 to 80 Aphis gossypii attacked per day. 

Predator larvae usually drop to the soil to pupate. They construct small silk 

cocoons in the top few millimetres of soil, but occasionally cocoons may be 

spun on plants. Larvae pupate within a few days of cocoon construction and 

adults emerge after 1Ð3 weeks depending upon temperatures. The life cycle 

can be completed in about 3 weeks at temperatures above 21¡C (Harris 1973; 

Herpai 1991). 

Harris (1973) does not record it from A. craccivora. Two 

hyperparasitoids of A. aphidimyza are known in Africa, the platygasterid 

Synopeas rhanis and an unidentified braconid (Harris 1973). 

There are many reports in the literature that natural enemies play an 

important role in reducing (and probably regulating) the abundance of pest 

aphids. More than 100 biological control programs have been mounted 

against at least 26 aphid species and 48% of them have reported success 

(HŒgvar and Hofsvang 1991). Twenty three species of aphidiid parasitoids 

have been used in classical biological control of aphids and the parasitoids 

became established in 32 out of 55 attempts (Greathead 1989). Most pest 

aphids are attacked in their native range by many parasitoids and predators 

and by a few pathogenic fungi. However, many of the natural enemies have 

not accompanied their aphid hosts when these have spread into new regions. 

Indeed, they may not even be present throughout the presumed native range 

of their host. Since both the direct and indirect damage caused by aphids 

seem to be proportional to their numbers, any reduction is potentially 

beneficial. Even in the case of virus transmission, where the feeding (or


probing) of single infected aphids on a crop may lead to substantial loss, 

reduction in aphid numbers will more than proportionally reduce the 

probability of flying aphids migrating to an uninfected plant (Wellings 

1991), because reduced crowding of aphids usually results in a lower 

number forming wings. As indicated above, there have been a number of 

attempts at classical biological control of aphids, and there are at least 7 

well-documented successes up to 1988 (Hughes 1989). None of these, 

however, involved A. craccivora or A. gossypii as the main target. These 

latter species have, however, been subjected to important attack by 

parasitoids introduced primarily against another pest aphid in the same 

general environment. For example, although Aphidius colemani failed to 

establish on Pentalonia nigronervosa in Tonga, it did so very successfully 

there on A. gossypii. Although A. craccivora is also present in Tonga, 

Aphidius colemani has not yet been recorded from it (Carver et al. 1993), but 

monitoring has been minimal. 

The best predictor of success in biological control is previous success 

with a natural enemy in a similar environment. If this experience is 

unavailable, the best chances appear to be with a climatically adapted, 

adequately host-specific, natural enemy that is known to attack the pest in its 

native or expanded range. Parasitic wasps appear to be the best natural 

enemies available for aphids, because they are generally far more host 

specific than predators and are often more efficient at searching for hosts at 

low aphid densities (Hughes 1989). Predators can be very effective in 

reducing aphid numbers at certain times of the year, but are often unable to 

prevent damage. Furthermore, their general lack of host specificity makes 

most of them unattractive to authorities responsible for approving 

introductions, so they have not been dealt with in any detail in this dossier. 

A feature that makes it difficult to generaliseÑand even to make clear 

recommendationsÑis that many of the identifications of some of the 

parasitoids are incorrect, particularly the earlier ones, but even some of the 

more recent ones made by non-specialists. The selection of appropriate 

natural enemies for an aphid biological control program requires a detailed 

knowledge of the ecology of the target aphid and often of other potential 

aphid hosts in the environment where it is causing problems; also of where to 

obtain parasitoid biotypes with appropriate host specificity and habitat 

adaptation. If an apparently good species fails to establish or, if established, 

to become effective, it is probably worth seeking the same enemy from a 

potentially more appropriate source, such as one with a better climate match; 

or a biotype that exhibits a special preference for the target aphid; or the first 

generation from field-collected material, rather than employing individuals 

bred for many generations in the laboratory (Hughes 1989).


Both A. craccivora and A. gossypii are now almost cosmopolitan in their 

occurrence throughout the temperate, subtropical and tropical regions of the 

world and both are polyphagous. At least A. gossypii exists as a series of 

biotypes with different spectra of host preferences and both it and 

A. craccivora owe a considerable amount of their economic importance to 

their ability to transmit an extensive range of important plant viruses. 

Both aphids are attacked by a wide range of parasitoids and share a 

number of these species. The majority of these parasitoids are polyphagous 

and attack many other (but not all) species of the genus Aphis and some 

species in related aphid genera. Two important parasitoids are the American 

Lysiphlebus testaceipes and the Indian Aphidius colemani. These parasitoids 

both exist in a number of biotypes with different host spectra and abilities to 

attack A. craccivora and A. gossypii on some plants, but not on others. Thus, 

when biological control is attempted, care must be taken to select a 

parasitoid biotype that is appropriate for the aphid biotype, the host plant and 

the prevailing environmental conditions. Laboratory comparison of the 

impact on A. gossypii of Aphidius colemani, Lysiphlebus testaceipes and 

Aphidius matricariae indicated that A. colemani was the most and 

A. matricariae the least effective (van Steenis 1992). The polyphagous 

nature of many effective parasitoids has the advantage that, in any region, a 

number of other aphid species will be parasitisedÑand hence serve as 

valuable reservoirs of parasitoids when the target pest population falls to a 

low level. The aims of aphid biological control are (i) as far as possible to 

maintain densities below those at which alates are formed due to crowding 

and (ii) if possible, to depress densities still further, so that sap removal, 

volume of honeydew produced and plant deformation become unimportant. 

Takada (1992) points out that the aphid parasitoid fauna in Far East Asia 

is quite different from that in India. Thus, the most important parasitoids of 

A. gossypii in the Far East are Trioxys communis, Lysiphlebia japonica and 

Aphelinus sp., none of which occurs in India. On the other hand, the 

dominant parasitoid of A. gossypii in India is Trioxys indicus, which is 

present in Taiwan, but not in Japan or Korea. Another Oriental species, 

Lipolexis scutellaris occurs in Hong Kong, Vietnam and Malaysia. It 

appears that Indian parasitoids, rather than Far East Asian species are 

present in Southeast Asia. However inadequate information is available in 

Southeast Asia on the natural enemies of A. craccivora and A. gossypii 

present in the many different crop systems and environments in which these 

aphids occur. Even within a single country, it is necessary to examine the 

aphid population in the particular situations and the crops where they are 

causing important problems. This can be illustrated by the parasitoid 

complex of A. gossypii in Japan where it occurs in habitats ranging from


open fields with low vegetation to garden habitats with low shrubs. In both 

situations it is attacked by specialised and generalist parasitoids. Of the two 

specialised parasitoids, Trioxys communis prefers the open field whereas 

Lysiphlebia japonica the garden habitat. Of the generalist parasitoids 

Aphelinus sp. prefers the open field and Ephedrus sp. the garden habitat. 

Thus the parasitoid complex of A. gossypii on cucumber, eggplant or taro in 

an open field is quite different from that on Hibiscus or Rhamnus in a garden 

(Takada 1992). Hence, in any country, the aim would be to determine 

whether there are gaps in the range of parasitoids present that might be filled 

with species known to be effective elsewhere. If there appear to be important 

gaps, there are good reasons for believing that there could be considerable 

advantages in filling them. Nevertheless, there are greater complexities than 

for many other pests in making clear recommendations, largely because of 

the range of biotypes that exist in both aphid hosts and their parasitoids. 

Although it is not possible to make specific recommendations for any 

country without knowing what parasitoids are already present and the 

crop(s) on which control is desired the following parasitoids merit 

consideration: 

Aphelinus gossypii Lysiphlebia japonica 

Aphidius colemani Lysiphlebus fabarum 

Aphidius gifuensis Lysiphlebus testaceipes 

Ephedrus persicae Trioxys communis 

Lipolexis scutellaris Trioxys indicus 

Even if any of these species is already in a country, but not attacking 

either A. craccivora or A. gossypii, a host-adapted strain should be 

considered for introduction. It is possible that some of the parasitoid species 

will compete directly with species that are already present. If this happens 

and one parasitoid species is displaced from an aphid host in some situations 

or on some crops, the final result will almost always be a lower overall aphid 

density.

4.5 Cosmopolites sordidus 

India 

20° 

Myanmar 

? Laos 

0° 

20° 

China 

P 

Thailand 

+ 

Cambodia 

+ 

Vietnam 

+++ 

++ 

++ Brunei 

Malaysia 

+ 

Singapore 

++ 

Indonesia 

+ 

Taiwan 

++ 

Philippines 

Australia 

Papua 

New Guinea 

++ 

The banana weevil borer Cosmopolites sordidus is native to the Indo-Malaysian region. 

There have been many attempts at biological control, involving three predatory beetles, but 

the results have generally been disappointing. Many predators attack Cosmopolites larvae in 

their native range, especially the histerid beetles Plaesius javanus and Plaesius laevigatus in 

Indonesia and Dactylosternum hydrophiloides in Malaysia. The first two species have been 

established in Fiji, with some reduction in Cosmopolites abundance, but it remains an important 

pest there. In Cook Is, they appear to have had little impact. Unless beneficial effects from P. 

javanus and P. laevigatus can be demonstrated in Fiji or other countries, there would seem to be 

little point in introducing these species to any additional countries. In Kenya, Koppenhšfer et al. 

(1992) recorded two important predatory beetles, Dactylosternum abdominale and 

Thyreocephalus interocularis, 

capable of reducing larval abundance by 40% to 90%. In Cuba, 

the ant Tetramorium bicarinatum is reported to keep C. sordidus under control. This ant is 

widespread, but there is no information on its effectiveness elsewhere. 

It would be highly desirable to investigate whether the weevil is indeed absent or of very 

minor importance in some areas (e.g. Myanmar, southern China) and, if so, what part is played 

by resistant cultivars, cultural methods or natural enemies. Ants might be evaluated for their 

effects in the Solomon Is where Cosmopolites is unimportant or in Papua New Guinea where it is 

of local importance only. 

85 

20° 

0° 

20°


Cosmopolites sordidus (Germar) 

Rating 

Origin 

Distribution 

Coleoptera: Curculionidae 

banana weevil borer 


+++ Viet +++ Cook Is, Fr. P, Fiji, Guam, 

New Cal, Niue, A Sam, Tong 

13 ++ Msia, Brun, Indo, 

Phil 

+ Thai, Sing + FSM 

P Camb P P Kir 

? Myan, Laos 

35 ++ PNG, Sol Is, Sam, Van, 

W & F 

This account brings up-to-date the chapter on C. sordidus in Waterhouse and 

Norris (1987) and increases its relevance to Southeast Asia. 

According to Purseglove (1972) the genus Musa has a centre of diversity in 

the Assam-Burma-Thailand area and it probably originated there. The 

banana weevil borer is also stated to be a native of the Indo-Malaysian region 

(Zimmerman 1968; Clausen 1978). Although this region seems a likely 

centre of origin of the weevil, and those investigating its biological control 

have consistently assumed so, there had already been, by the time of 

GermarÕs 1824 description based on material from India, centuries of 

intercontinental travel by Europeans, by means of which the weevil could 

have spread to many other lands in infested plants, thus obscuring its origin. 

There are only two species in the genus Cosmopolites, 

the lesser known 

C. pruinosus occurring in Borneo and the Philippines and, after introduction, 

in Micronesia (Zimmerman 1968). 

C. sordidus is present in virtually all banana-growing areas of the world, 

including most, if not all, of Southeast Asia and most of the Pacific. 

Exceptions in the Pacific are Marshall Is, Tuvalu and Tokelau (Anon. 1979b; 

Waterhouse and Norris 1987; Waterhouse 1997). In Southeast Asia no 

information is available from Laos and the situation in Myanmar is unclear. 

A recommendation was made in the standard work on the ÔInsect Pests of 

BurmaÕ (Ghosh 1940) to guard against the introduction to that country of 

C. sordidus and neither N. von Keyserlingk nor G. Pierrard (pers. comm.

Biology 

4.5 


1992) were able to establish, when based in Yangon, that the species 

occurred in Myanmar. Its uncertain status in Myanmar and Laos and its 

status of Ôpresent, but unimportantÕ, in Cambodia and China clearly merits 

further investigation. 

The ovoid, 2 mm long, white eggs are laid singly, usually between the leafsheath 

scars on the crown of the banana rhizome, in small cavities chewed 

out by the female just above the ground surface. Laying also occurs on the 

pseudostems of fallen plants. The eggs hatch in about 8 days in summer and 

the larvae tunnel into the tissues. On reaching maturity, after about 20 days 

feeding in warm weather (Jepson 1914), the larvae tunnel to near the surface 

of the corm and form an oval chamber in which they pupate. The period from 

egg to adult may be as short as 29 days in the New South Wales summer or as 

long as 6 months in the cooler parts of the year (Hely et al. 1982). Fifty days 

is a more usual maximum for the life cycle in Fiji (Swaine 1971). The 

nocturnal adults also tunnel in banana tissues. During the day they generally 

hide in or around the rhizomes or between the leaf sheaths at or just above 

ground level. They are slow moving and will feign death when disturbed. 

They seldom fly, but walk over the soil surface and vegetation. Whalley 

(1957) found that adults dispersed slowly in Uganda. Of 400 marked weevils 

35% were recovered over an 8-month period within a radius of 7 m from the 

release point. In Colombia a few marked adults were recaptured after 2 

weeks at a distance of 6 to 8 m, but 23 months after release none were found 

in another plantation 20 m away (Cardenas and Arango 1986). Male 

C. sordidus release an aggregation pheromone, sordidin (Beauhaire et al. 

1995), from the hindgut which attracts both males and females, but females 

do not produce a pheromone attractive to either sex (Budenberg et al. 1993b; 

Ndiege et al. 1996). Both sexes were attracted to freshly cut rhizome and 

pseudostem and females to rotting pseudostem (Budenberg et al. 1993a). 

Eggs are laid throughout the year at a rate varying with temperature and up to 

100 a year. Adults may live as long as 2 years (Swaine 1971). They can 

survive in captivity for 14 weeks without food (Zimmerman 1968). 

Laboratory rearing of C. sordidus was studied by Afreh (1993) and 

Koppenhšfer and Reddy (1994). 

87


Damage 

The status of C. sordidus as one of the most important pests of bananas is 

often reported (Swaine 1971; Purseglove 1972) and, indeed, many adults 

and larvae are often present. It is reported to be now the most important pest 

of bananas in Africa (Nahif et al. 1994; Ortiz et al. 1995). However, in order 

to place these reports in context, it is necessary to outline the stages of 

growth of the plant. Bananas are propagated vegetatively by planting 

rhizomes (corms), which give rise to shoots after a few weeks. As the plant 

grows, a pseudostem is formed from the sheaths of the leaves which 

continue to develop internally until a flowering shoot emerges from the top 

of the pseudostem at a height of 2 to 4 m depending upon the variety. When 

each bunch of bananas is cut, the pseudostem bearing that bunch is also cut 

down. At the same time a healthy sucker growing from the same base is 

selected to succeed and other suckers removed. C. sordidus larvae tunnel in 

the rhizome and the base of the pseudostem, but do not attack the roots. This 

tunneling may kill young plants and greatly increases the susceptibility of 

mature plants to wind damage. Adults cause little damage and feed mainly 

on rotting banana tissue. Injury by larvae to the rhizome can interfere with 

root initiation and sap flow within the plant and grossly infested plants may 

bear small bunches of undersized fruit (Wright 1976). Much of the damage 

attributed to C. sordidus is probably caused by rhizome rot or nematodes 

(Ostmark 1974). In East Africa the combined attack of nematodes and of 

banana weevil borer is considered to be the main reason for the serious 

decline in productivity of bananas (De Langhe 1988), but the precise role of 

the weevil is still to be established. Suckers infested with nematodes were 

found to be more than four times more likely to be attacked by C. sordidus 

than suckers without nematodes (Speijer et al. 1993). Although the banana 

weevil borer has occasionally been responsible for severe losses of newlyplanted 

rhizomes, extensive experiments in Central America agree with 

some reports from Australia (Smith 1993; Wallace 1937) that weevil 

damage is not as important as frequently claimed since the larvae have a 

strong preference for rhizomes of harvested plants over healthy rhizomes 

(Ostmark 1974). There are reports that some banana cultivars are 

comparatively resistant to attack by C. sordidus (Mesquita et al. 1984; 

Mesquita and Caldas 1987), but the mechanism of such resistance 

(repellency, toxicity, greater tolerance to damage, etc.) is not known. 

In Australia, Braithwaite (1963) concluded that the importance of 

C. sordidus infestation is aggravated by poor culture, but that benefit could 

be derived from almost complete control with insecticides (Braithwaite 

1958). In the same region Loebel (1975) concluded that heavy weevil

Host plants 

4.5 


infestation is a symptom, rather than a cause, of a declining plantation, 

because 2 years of effective use of chemicals failed to improve growth or 

yield in his experimental plots. In Costa Rica several insecticides were 

effective in controlling C. sordidus populations, but banana yields were not 

increased (Nanne and Klink 1975). Nevertheless C. sordidus is always likely 

to be of importance in areas that experience strong winds. The abundance of 

adult weevils can be estimated by counting the number attracted to cut 

segments of pseudostem and of larvae by estimating the area damaged and 

counting the number of galleries exposed by slitting the rhizome or the 

pseudostem very near to its base (Vilardebo 1973; Delattre 1980; Mesquita 

1985; Smith 1993). In spite of this, an adequate relationship between adult 

and larval abundance and economic loss remains to be established. It must 

be added, however, that there continues to be a widespread view that 

C. sordidus is a major pest. 

The weevil attacks all banana ( Musa sapientum) 

cultivars, including 

plantain, and also Manila hemp ( Musa textilis). 

It has been recorded in 

earlier days (but not in recent years) from plants in other Orders, but these 

reports are almost certainly in error. 


Although many predators are known to attack C. sordidus eggs, larvae and 

pupae (Table 4.5.1), it is extraordinary that, with one possible exception, not 

a single parasitoid of any life history stage has been recorded. That exception 

is the early report from the Philippines of Cendana (1922), who found a 

chalcidid wasp in one of his C. sordidus breeding cages, but it may not have 

been parasitising the weevil. It is, perhaps, less surprising that the heavily 

sclerotised adult weevil has very few enemies. It is true that some weevils 

are effectively attacked by hymenopterous parasitoids and that some 

tachinid parasitoids are able to attack certain weevils by laying eggs in their 

food or beneath their mouth when they are feeding (Jacobs and Renner 

1988), but neither has been observed for C. sordidus. 

Koppenhšfer 

(1993a,b) estimated that 58% of the eggs were accessible to predators and 

presumably at least as many should be available to parasitoids if there were 

any. Most eggs were found in the surface layer of rhizomes, particularly in 

the crown, but some are also laid at the base of pseudostems and in the walls 

of abandoned larval tunnels in both pseudostems and rhizomes. As soon as 

eggs hatch the young larvae immediately tunnel deeper into the plant tissue 

and thus become far less available to natural enemies. 

89

Table 4.5.1 

Insect predators of Cosmopolites sordidus 

Insect 

DERMAPTERA 

LABIIDAE 

Stage 

attacked 

Carcinophora (= Psalis) 

americana 

larvae Jamaica 

Puerto Rico 

Euborellia (= Anisolabis) 

annulipes 

egg, larva Jamaica 

Kenya 


Edwards 1934 

Anon. 1939 

Edwards 1934 

Koppenhšfer et al. 1992; Sirjusingh 

et al. 1992 

Labia borellii 

egg, larva Kenya Koppenhšfer et al. 1992 

Labia curvicauda 

HEMIPTERA 

CYDNIDAE 

egg, larva Kenya Koppenhšfer et al. 1992 

Geotomus pygmaeus 

MIRIDAE 

egg Malaysia, widespread China 1935 

Fulvius nigricornis 

NABIDAE 

egg Malaysia China 1935 

Phorticus pygmaeus 

REDUVIIDAE 

egg Malaysia, Papua New Guinea China 1935 

Physoderes curculionis 

COLEOPTERA 

CARABIDAE 

larva Malaysia China 1935 

Abacetus? 

optimus 

Kenya Koppenhšfer et al. 1992 

Galerita (= Propagalerita) 

bicolor 

Sirjusingh et al. 1992 

Scarites sp. Sirjusingh et al. 1992 


Table 4.5.1 (contÕd) Insect predators of Cosmopolites sordidus 

Insect Stage 

attacked 

COLEOPTERA 

ELATERIDAE 

unidentified spp. Australia, 

New Caledonia 

HISTERIDAE 

Hister niloticus 

Hololepta (= Lioderma) 

quadridentata 


Froggatt 1928a 

Jacques 1931 

larva Kenya Koppenhšfer et al. 1992 

Malaysia 

Trinidad 

Clement 1944 

Pea & Duncan 1991 

Hololepta striaditera 

larva, pupa Kenya Koppenhšfer et al. 1992 

Hololepta sp. St Vincent Sirjusingh et al. 1992 

Lioderma sp. Sirjusingh et al. 1992 

Plaesius (= Hyposolenus) 

laevigatus 

Indonesia Froggatt 1928b 

Plaesius javanus 

larva, pupa Malaysia 

Froggatt 1928b; Clement 1944; 

Indonesia 

Jepson 1914 

Thailand 

Charernsom 1992 

Platysoma abrupta 

larva Malaysia Corbett 1936; Lamas 1947 

Platysoma sp. Corbett 1936 

unidentified histerid sp. 

HYDROPHILIDAE 

egg, larva, pupa Kenya Koppenhšfer et al. 1992 

Dactylosternum abdominale 

larva Kenya, 

Corbett 1936; Edwards 1939, 

Philippines 

Koppenhšfer et al. 1992 

D. hydrophiloides larva Indonesia, 

Malaysia 

Corbett 1936 

D. intermedium larva Guinea CuillŽ 1950 

D. profundus San Thom Beccari 1967 

D. subdepressum Trinidad Cock 1985 

D. subquadratum larva Malaysia Corbett 1936 

Omicrogiton insularis larva Malaysia Corbett 1936 

4 

.5 


91

Table 4.5.1 (contÕd) Insect predators of Cosmopolites sordidus 

Insect Stage 

attacked 

COLEOPTERA 

SILVANIDAE 

Cathartus sp. larva Indonesia Jepson 1914 

STAPHYLINIDAE 

Belonuchus ferrugatus larva Indonesia Jepson 1914; CuillŽ 1950 

B. quadratus larva Malaysia Corbett 1936; Lamas 1947 

Charichirus sp. egg. larva Kenya Koppenhšfer et al. 1992 

Eulissus sp. Kenya Reddy 1988 

Hesperus? sparsior egg, larva Kenya Koppenhšfer et al. 1992 

Priochirus (= Leptochirus) unicolor larva Indonesia Jepson 1914, CuillŽ 1950 

Thyreocephalus? interocularis egg, larva Kenya Koppenhšfer et al. 1992 

TENEBRIONIDAE 

Eutochia pulla egg Kenya Koppenhšfer et al. 1992 

DIPTERA 

RHAGIONIDAE 

Chrysopilus ferruginosus larva Indonesia, 

India, 

Philippines 


Froggatt 1928b; Beccari 1967 

Jepson 1914 

Chrysopilus sp. Brazil Sirjusingh et al. 1992 

HYMENOPTERA 

FORMICIDAE 

Anochaetus sp. Kenya Reddy 1988 

Pheidole megacephala egg, larva Cuba Castineiras et al. 1991a 

Tetramorium bicarinatum 

(= T. guineense) 

egg, larva Cuba Roche & Abreu 1983 


4.5 Cosmopolites sordidus 93 

Because of the reported existence of natural enemies of C. sordidus in 

Indonesia and nearby countries, Jepson (1914) was sent from Fiji to 

investigate the situation in Java. The histerid beetle Plaesius javanus was 

commonly found preying on C. sordidus and other insects in the soil and leaf 

litter. Two staphylinid beetles Belonuchus ferrugatus and Priochirus 

(= Leptochirus) unicolor and a silvanid beetle Cathartus sp. were also 

shown to attack C. sordidus larvae, but they were not nearly as voracious. 

The predatory larvae of the rhagionid fly Chrysopilus ferruginosus attacked 

C. sordidus larvae in the laboratory, but not in the field. 

A later investigation in Java (Froggatt 1928b) failed to reveal any egg 

parasites, but P. javanus and C. ferruginosus were recorded, as well as two 

other species of Histeridae, one probably Plaesius (= Hyposolenus) 

laevigatus, one or two species of Staphylinidae and two species of 

Hydrophilidae (all Coleoptera). Several species of Dermaptera were fairly 

common in the rotting banana plant. In southern China (Yunnan Province) 

two Dermaptera (one a forficulid) are reported to eat C. sordidus larvae and 

pupae, and a mite to attack larvae and adults; also a white muscardine fungus 

to infect 1% of larvae and pupae (Sun 1994). 

The only detailed study in recent times of the natural enemies of 

C. sordidus is that of Koppenhšfer et al. (1992) in Kenya. Not surprisingly, 

the species (of predators) recorded were all polyphagous, since many were 

native species that had come to include an introduced pest amongst their 

prey. Twelve predators of eggs, larvae and pupae of C. sordidus were found. 

None of these attacked adults and no parasitoids were recorded 

(Koppenhšfer et al. 1992). Of these predators, the hydrophilid beetle 

Dactylosternum abdominale reduced weevil numbers in suckers by up to 

50% and in residual stumps of harvested suckers by 39%. In spent 

pseudostems, D. abdominale reduced numbers by 40% to 90% at different 

predator densities and the large staphylinid predator Thyreocephalus 

interocularis reduced numbers by 42%. Other predators (Table 4.5.1) were 

unimportant (Koppenhšfer and Schmutterer 1993). 

These particular predators are clearly non-specific, since they are native 

to East Africa and C. sordidus is not. Thus, although they may be attractive 

candidates for introduction (and Plaesius javanus was in this same 

category), the widely held view now is that, because of their impact on nontarget 

organisms, very careful consideration should be given before any 

highly non-specific predators are introduced to a new region. 

Extensive testing has been carried out of fungi (Metarhizium anisopliae 

and Beauveria bassiana) and of entomopathogenic nematodes 

(Heterorhabditis spp. and Steinernema spp.) as biological ÔpesticidesÕ 

against C. sordidus. Laboratory or field trials showed that strains of both


fungi were capable of killing adults and larvae (Delattre and Jean-Bart 1978; 

Filho et al. 1987; Busoli et al. 1989; Castineiras et al. 1991a,b; Pea and 

Duncan 1991; Ponce et al. 1992; Brenes and Carballo 1994; Carballo and de 

Lopez 1994; Pea et al. 1995). In the field the best results were obtained with 

application of fungi twice a year at a dose of 10 13 conidia per ha. This 

reduced trap catches of adults by 52% and rhizome damage by 65%, leading 

to a 25% yield increase. Parallel experiments with 9 colonies per ha of the 

predatory ant Pheidole megacephala yielded similar results (Castineiras et 

al. 1991a,b). 

Early laboratory tests in Guadelupe by Laumond et al. (1979) showed 

that C. sordidus is susceptible to the entomopathogenic nematode 

Steinernema carpocapsae (= S. feltiae), an observation since widely 

confirmed in Central America for this and other nematode species (e.g. 

Figueroa 1990; Pea and Duncan 1991). However, the most extensive recent 

work has been carried out in Australia and Tonga. Twenty-one different 

species of Steinernema and Heterorhabditis, 7 strains of Steinernema 

carpocapsae, 2 of S. feltiae, 4 of H. bacteriophora and 2 of 

H. zealandica were tested against adult banana weevils. The best of these, 

S. carpocapsae BW strain, gave 85% infection in the laboratory (Parnitzki et 

al. 1990, 1998; Treverrow et al. 1991). Adult C. sordidus are highly resistant 

to entomopathogenic nematodes, due to the difficulty of nematodes entering 

the host via the mouth or anus. The large spiracles of the first abdominal 

segment offer an effective site of entry for the nematodes if they are able to 

pass under the tightly fitting elytra. By adding paraffin oil to the nematode 

preparation to seal the elytra, the beetle is caused to raise them slightly to 

respire, simultaneously giving the nematodes access to the spiracles. Adult 

weevils are strongly attracted to holes or cuts in the rhizome or psuedostem, 

but they require a thigmotactic stimulus to remain long enough to become 

infected by nematodes. If a core of tissue is removed from two sites at the 

base of a residual corm using a desuckering tool, 250000 nematodes 

(S. carpocapsae BW) added to each hole and the core loosely inserted, 

nearly all adult weevils attracted are killed. In one large scale field trial in 

New South Wales 8% of plants in untreated plots suffered significant 

damage, 3% when prothiophos was added to the core, 1% when nematodes 

were added and 0% when prothiophos was applied to the soil around the base 

of the plant. It was concluded that control of banana weevil using 

entomopathogenic nematodes should now be economically feasible 

(Treverrow and Bedding 1992, 1993a,b). More recently, Treverrow (1994) 

has found that baiting, and stem injection with very small amounts of 

insecticide, can reduce treatment costs to less than 1 cent per stool. This 

makes nematode applications against adults uncompetitive unless market


premiums can be obtained for fruit produced in the absence of insecticides. 

However, targetting the highly susceptible larvae instead of the more 

resistant adult weevils may significantly reduce the costs of nematode 

applications. Applications of S. carpocapsae with a water-absorbing 

polyacrylamide gel into cuts or holes made in residual rhizomes gave 

significant mortality of C. sordidus larvae (Treverrow et al. 1991; Treverrow 

and Bedding 1993b). However, mortality at this stage may have limited 

effect on abundance, since a survey of 50 properties showed that 70% of 

adult weevils had already emerged from pre-harvest corms (Treverrow and 

Bedding 1993a; Treverrow 1994). The importance of correct formulation for 

the nematodes is highlighted by the disappointing results obtained in 

Queensland, when an earlier formulation without a water-absorbing gel 

failed to give effective control of adults, possibly due to the accumulation of 

excess water in the core holes (Smith 1993). 


FIJI 

C. sordidus became a very destructive pest of bananas following its 

introduction about 1901 and this led to the first attempt at its biological 

control. This consisted of the introduction of the predatory histerid beetle 

Plaesius javanus into Fiji in 1913 from Java (Jepson 1914). This was 

unsuccessful (Table 4.5.2), but a further introduction in 1918 led to its 

establishment (Veitch 1926; Bennett et al. 1976). Simmonds (1935) reported 

a marked reduction in weevil damage, an opinion supported by Anon. 

(1935), but Pemberton (1954) considered that only partial control had been 

achieved. It now seems probable that, in addition to P. javanus, the similar 

looking histerid Plaesius laevigatus was also introduced, at least to Cook Is 

(Walker and Deitz 1979). 

AUSTRALIA 

Cosmopolites sordidus is thought to have become established in Queensland 

about the end of the 19th century, having arrived possibly from Papua New 

Guinea, but it was not until about 1914Ð15 that it reached New South Wales 

as a result of an accidental introduction from Fiji (Wilson 1960). Following 

its successful introduction into Fiji, Plaesius javanus was introduced from 

Java and liberated in Queensland from 1921 to 1928, but became established 

only briefly. It was introduced into New South Wales in 1922 from Java and 

again in 1934 from Fiji, but again it did not become established. The fly 

Chrysopilus ferruginosus was also introduced from Java, but failed to 

become established.

Table 4.5.2 Introductions for the biological control of Cosmopolites sordidus 

Country and species Liberated From Result Reference 

AUSTRALIA 

Plaesius javanus 1921Ð28 

Java 

Ð Clausen 1978 

Weddell 1932 

1934 

Fiji 

Ð Wilson 1960 

Chrysopilus ferruginosus 1928 Java Ð Wilson 1960 

Dactylosternum hydrophiloides 

CAMEROON 

1939 Malaysia + Smith 1944; Wilson 1960 

Plaesius javanus 1952 Trinidad Ð Bennett et al. 1976 

Hololepta quadridentata 1952 Trinidad Ð Bennett et al. 1976 

COOK IS 

Plaesius javanus 1937Ð40 Fiji + Walker & Deitz 1979 

Plaesius laevigatus 1937Ð40 Fiji + Walker & Deitz 1979 

CUBA 

Plaesius laevigatus Ð Sirjusingh et al. 1992 

DOMINICA 

Plaesius javanus 1951 Trinidad Ð Cock 1985 

1958Ð59 ? Ð Clausen 1978 

Hololepta quadridentata 1951 

1958Ð59 

FIJI 

Trinidad 

? 

Ð 

Ð 

Cock 1985 

Clausen 1978 

Plaesius javanus 1913 Java Ð Jepson 1914 

1918 Java + Veitch 1926 

Plaesius laevigatus 1918 Java + Walker & Deitz 1979 


Table 4.5.2 (contÕd) Introductions for the biological control of Cosmopolites sordidus 

Country and species 

FRENCH POLYNESIA 

Liberated From Result Reference 

Plaesius javanus 1937 Fiji + Delobel 1977 

GRENADA 

Hololepta quadridentata 1949, 1951 Trinidad Ð Cock 1985 

Plaesius javanus 1949, 1951 Trinidad Ð Cock 1985 

HONDURAS 

Plaesius javanus 1942 ? Ð Greathead 1971 

JAMAICA 

Plaesius javanus 1918Ð19 Java Ð Cock 1985; Edwards 1934 

1937Ð38 Java via Fiji + 

Ð 

Cock 1985 

Sirjusingh et al. 1992 

Dactylosternum hydrophiloides 1918Ð19 Malaysia Ð Edwards 1934 

1937Ð38 Malaysia + Edwards 1934 

Dactylosternum abdominale 1937Ð38 Malaysia Ð Edwards 1934 

MARIANAS 

Plaesius javanus 1947 Fiji + Clausen 1978 

Hololepta quadridentata 1953 ? Ð Clausen 1978 

Hololepta minuta 1953 ? Ð Clausen 1978 

Hololepta sp. 1953 Trinidad Ð Clausen 1978 

MAURITIUS 

Plaesius javanus 1959 Trinidad Ð Bennett et al. 1976 

? Trinidad Ð Gomy 1983 

Hololepta quadridentata 1942 Trinidad Ð Bennett et al. 1976 

1959 ? Ð Clausen 1978 

4.5 Cosmopolites sordidus 97



MEXICO 



NEW CALEDONIA 

1955 ? + Barrera & Jiminez 1994; 

Greathead 1971 


PALAU IS 

1949 Fiji + Dumbleton 1957 

Dactylosternum hydrophiloides 1948 Malaysia Ð Dumbleton 1957 

Hololepta sp. 1953 ? Ð Dumbleton 1957 

PUERTO RICO 


SAMOA 

Plaesius javanus 1957 Fiji + Dale and Herring 1959 

SEYCHELLES 

Plaesius javanus 1952 Trinidad Ð Greathead 1971 

Hololepta quadridentata 1952 Trinidad Ð Greathead 1971 

ST LUCIA 

Hololepta quadridentata 1950Ð1954 Trinidad Ð Cock 1985 

Plaesius javanus 1950Ð1954 Trinidad Ð Cock 1985 

ST VINCENT 

Dactylosternum subdepressum 1950Ð1954 Trinidad Ð Cock 1985 

Hololepta quadridentata 1942 Trinidad + Bennett et al. 1976 

Plaesius javanus 1981 Trinidad Ð Cock 1985 




TAIWAN 



TANZANIA 


TONGA 

Plaesius javanus 1952 Fiji Ð Dumbleton 1957 

TRINIDAD 

+ O. Fakalata pers. comm. 

Plaesius javanus 1942 Jamaica + Bennett et al. 1976 

UGANDA 

Ð Sirjusingh et al. 1992 


VANUATU 

1934Ð35 Java Ð Greathead 1971 

Plaesius javanus ? ? ? Anon. 1979a 

WALLIS IS 

Plaesius javanus 1947 Fiji ? Cohic 1959 

4.5 Cosmopolites sordidus 99


The predatory hydrophilid beetle Dactylosternum hydrophiloides from 

Malaysia was liberated in 1939 and has become established but has not had a 

major effect on weevil abundance (Wilson 1960). 

Braithwaite (1958) reports an unusual native predator of C. sordidus, a 

blue planarian worm Kontikia (= Geoplana) caerulea, which lives in moist 

sheltered situations. It sucks out the body fluids of its prey, leaving the 

cuticle intact. 

CAMEROON, MAURITIUS, SEYCHELLES, UGANDA 

Both P. javanus and Hololepta quadridentata were supplied from Trinidad 

to Cameroon (1952), Mauritius (1959) and the Seychelles (1950Ð54) and 

P. javanus from Java to Uganda in 1934Ð35. Neither species became 

established (Greathead 1971). 

COOK IS 

CUBA 

INDIA 

JAMAICA 

Although only Plaesius javanus is recorded as having been introduced from 

Fiji into the Cook Is during the period 1937 to 1940, voucher specimens 

(DSIR, NZ) show that Plaesius laevigatus was also present in the material 

liberated and both species still occur in the Cook Is, although the latter does 

not appear in voucher specimens in Fiji. Unfortunately the banana weevil 

borer is still an important problem (Walker and Deitz 1979; Waterhouse 

1995, 1997). 

In Cuba the ant Tetramorium bicarinatum (= T. guineense) was found to be 

capable of destroying up to 65% of C. sordidus in heavily infested 

plantations. With lower populations, 73.8% and 83.5% control was obtained 

in successive years. For colonisation of a plantation in 3 to 4 months, ants 

should be released over 25% to 30% of the area (Roche and Abreu 1983). 

The ant is a more effective predator during the dry than the wet season 

(Lopez and Ramos 1986). In countries where this widespread ant is already 

present (e.g. the Americas, Africa, Papua New Guinea, Australia, the 

oceanic Pacific) it might well be considered for manipulating C. sordidus 

abundance, but very careful consideration should be given to any proposal to 

introduce such an agressive broad-spectrum predator into a new country. 

The predatory beetle Dactylosternum hydrophiloides was introduced from 

Malaysia in 1948, but there is no record of the outcome (Whilshaw 1949). 

Although an initial release in 1918Ð19 of Plaesius javanus from Java was 

unsuccessful, this predator became established as a result of a further release 

of Fijian material in 1937Ð38 (Bennett et al. 1976). More recently Sirjusingh 

et al. (1992) recorded that it was no longer present.


MYANMAR 

There do not appear to be any records in Myanmar of natural enemies of 

C. sordidus which is very uncommon and apparently confined to aromatic 

and sweet-flavoured banana varieties. Control is achieved by cutting the 

pseudostems every three years, a practice readily accepted because the stems 

are used in Mohinga, a popular fish soup (H. Morris pers. comm. 1994). 

PAPUA NEW GUINEA 

Bananas are grown widely and are the staple food in some areas but 

C. sordidus is not a serious pest. The ant Tetramorium bicarinatum occurs 

there but there is no information on any possible interaction with C. sordidus 

(J.W. Ismay pers. comm. 1985). 

TRINIDAD 

P. javanus was established in Trinidad in 1942 and, from there, together with 

a native histerid Hololepta quadridentata, it was sent to other islands in the 

West Indies. Evidence of establishment was not available to Simmonds 

(1958) but, in 1972, H. quadridentata was recovered in St Vincent (Bennett 

et al. 1976). 

OTHER COUNTRIES 

Although details are not available, P. javanus has been widely distributed 

and is reported to be established in French Polynesia, Marianas, New 

Caledonia (where chemical control is still required) (Delobel 1977; Clausen 

1978; M. Kauma pers. comm. 1985) and Tonga (O. Fakalata pers. comm. 

1985). It is also widespread on Upolu Is, Samoa (T.V. Bourke pers. comm. 

1986). 

In addition to the records in table 4.5.2, Sirjusingh et al. (1992) list a 

number of predators for Central or South America (Table 4.5.3). However, it 

is seldom clear which of these may be introductions (intentional or 

otherwise) from elsewhere and which are native to the region, as some 

almost certainly are. Their biological control potential has not been assessed.


Table 4.5.3 Additional natural enemies of C. sordidus in Central and 

South America 

Insect 

DERMAPTERA 

LABIIDAE 

Country ?Native 

Carcinophora (= Psalis) americana Brazil probably 

Euborellia annulipes 

HEMIPTERA 

Brazil ? 

CYDNIDAE 

Geotomus pygmaeus Brazil probably not 

MIRIDAE 

Fulvius nigricornis Brazil probably not 

NABIDAE 

Phorticus pygmaeus Brazil probably not 

REDUVIIDAE 

Physoderes curculionis Brazil probably not 

COLEOPTERA 

CARABIDAE 

Galerita bicolor Florida probably 

Scarites spp. 

HISTERIDAE 

Florida probably 

Hololepta spp. St Vincent possibly 

Lioderma sp. Brazil probably 

Platysoma abrupta 

HYDROPHILIDAE 

Brazil probably not 

Dactylosternum abdominale Trinidad probably not 

D. hydrophiloides Trinidad probably not 

D. intermedium Trinidad probably not 

D. profundus Trinidad probably 

Omicrogiton insularis 

SILVANIDAE 

Brazil probably not 

Cathartus sp. 

STAPHYLINIDAE 

Brazil (not established) 

Belonuchus ferrugatus Brazil (not established) 

B. quadratus Brazil probably not 

Priochirus (= Leptochirus) unicolor 

DIPTERA 

RHAGIONIDAE 

Brazil (not established) 

Chrysopilus sp. Brazil (not established)

Biology of main natural enemies 


Dactylosternum abdominale Col.: Hydrophilidae 

This is the most effective predator in Kenya. Its larvae are polyphagous 

predators and consume the contents of C. sordidus larvae and at high 

predator density may become cannibalistic. They also feed on the microfauna 

and micro-flora of decomposing plant tissues. On the other hand, the 

adults cause significant mortality of C. sordidus eggs; although many are 

laid in inaccessible positions inside the pseudostem and the polyphagous 

adults do not specifically search for eggs. Adults cannot penetrate the 

narrow tunnels of young larvae, so can only capture newly hatched larvae: 

later instar larvae are not attacked (Koppenhšfer and Schmutterer 1993; 

Koppenhšfer et al. 1992, 1995). Development from egg to adult takes 17 to 

33 days, life span is 95 days and females lay an average of 1.7 egg cases per 

week, each case containing 4 eggs. The preoviposition period is 16.6 days 

(Koppenhšfer et al. 1995). 

Geotomus pygmaeus Hem.: Cydnidae 

This predator is recorded from India, Ceylon, Myanmar, Indonesia, 

Vietnam, China, Japan, New Caledonia, Fiji, Samoa, French Polynesia and 

Hawaii. Its extensive distribution is probably due to its ready transportation 

in the soil attached to the roots of cultivated plants. Although reported to 

attack C. sordidus eggs in Malaysia, China (1935) suggests that this species 

normally is unlikely to be a predator. 

Plaesius javanus Col.: Histeridae 

The predatory larvae and adults of this Indonesian beetle attack larvae and 

pupae of Cosmopolites sordidus and many other soil and litter-inhabiting 

insects. Eggs are laid singly in old banana stumps, at the base of the stem and 

on the rhizome below the soil surface. The eggs hatch in 8 to 9 days, 

producing active, voracious larvae which feed for 5 to 6 months, older larvae 

being capable of consuming up to 30 or more C. sordidus larvae per day. A 

prepupal period of 10 to 15 days is passed in a pupation cell constructed in 

the soil, followed by a pupal stage of about 14 days. The adult remains in the 

cell for 7 to 10 days before emerging and is then capable of consuming 7 or 8 

weevil larvae per day (Clausen 1978; Jepson 1914; Weddell 1932). 

Thyreocephalus interocularis Col.: Staphylinidae 

This is the second most effective predator on C. sordidus larvae in Kenya. 

Both adults and larvae are polyphagous. In the absence of other hosts adults 

may prey on their own larvae and larvae are also occasionally cannibalistic 

(Koppenhšfer and Schmutterer 1993). After a pre-oviposition period of 32 

days, females lay an average of 31 eggs in decomposing banana


Comments 

pseudostems and in moist soil below banana mulch. Pupation occurs in the 

soil. Total development time averages 46 days and adults live an average of 

142 days (Koppenhšfer 1994). 

Although weevils, as a group, seem to be poor candidates for biological 

control, the establishment of Plaesius javanus and P. laevigatus in Fiji 

appears to have reduced the pest status of Cosmopolites sordidus there. 

Introductions of P. javanus (Table 4.5.2) have resulted in successful 

establishment (but often not at the first attempt) in Cook Is, French 

Polynesia, Jamaica, Marianas, Mauritius, New Caledonia, Samoa and 

Trinidad, but no information is available on the effects it has produced. 

Introductions have been unsuccessful in Australia, Cameroon, Dominica, 

Honduras, Mauritius, Mexico, Puerto Rico, Seychelles, St Lucia, St Vincent, 

Taiwan, Tanzania, Tonga and Uganda (Bartlett 1937; Miwa 1938; Clausen 

1978). Based on his observation and that of others, Koppenhšfer (1993a,b, 

Koppenhšfer and Schmutterer 1993) considered that the biology of 

P. javanus does not enable it to have any greater effect and that studies are 

necessary of the potential impact of proposed predator species before 

release. Two other predators have been established, one in Australia and one 

in Jamaica and St Vincent, but seemingly without much effect. 

If the beneficial effects of P. javanus (and P. laevigatus) in Fiji can be 

confirmed, it may be worth renewing efforts to establish them in other 

countries where C. sordidus is a major pest. Otherwise, resources available 

for biological control might be better deployed searching for, and 

evaluating, other natural enemies. 

It is possible that entomopathogenic nematodes (CSIRO 1993; 

Treverrow and Bedding 1993a) or fungi may hold some promise as 

biological pesticides. Nematodes generally have far less capability for selfperpetuation 

and dispersal in the environment than imported arthropod 

enemies, but they can be highly effective. However, in many tropical 

countries where bananas are a major staple food the distribution and 

application of mass-produced biological control agents, such as nematodes 

or fungi, is often impracticable because of storage and transport problems 

and lack of suitable equipment for application. In addition, like insecticides, 

they may be too costly for subsistence farmers who constitute the majority of 

banana producers. Even if it only leads to a partial (but still significant) 

reduction in pest status, classical biological control is, under these 

circumstances, a particularly appropriate approach to reduce losses.

4.6 Deanolis sublimbalis 

India 

20° 

0° 

20° 

Myanmar 

Laos 

China 

++ 

Thailand 

+ 

Cambodia 

Malaysia 

Vietnam 

Singapore 

P 

Brunei 

P 

Indonesia 

+ 

Taiwan 

++ 

Philippines 

Australia 

+ 

Papua 

New Guinea 

++ 

105 

The red banded mango caterpillar, Deanolis sublimbalis tunnels in the flesh and seed of the 

fruit of mango, Mangifera indica, 

and also attacks the fruit of M. odorata, 

M. minor and 

Bouea burmanica. 

It is reported from India eastwards to Southeast Asia, southern China 

and Papua New Guinea. In this vast region there are scattered reports of damage ranging 

up to 50 per cent of fruit, but many areas within it from which there are no reports of damage 

or even of its presence. 

The only records of natural enemies are of two trichogrammatid egg parasitoids and a 

vespid larval predator, all in the Philippines. Further searches would be necessary before 

the potential of classical biological control could be evaluated, especially for infestation in 

new areas into which it has spread in recent years. 

20° 

0° 

20°


Deanolis sublimbalis Snellen 

Synonyms 

Rating 

Origin 

Distribution 

Lepidoptera: Pyralidae: Odontinae 

red banded mango caterpillar, red banded borer 

Long known as Noorda albizonalis Hampson 1903 or Autocharis 

albizonalis (Hampson), this species should be referred to as Deanolis 

sublimbalis Snellen, because of the priority of its description (Snellen 1899) 

(M. Shaffer pers. comm. 1997) from specimens collected in Celebes (now 

Sulawesi). He also referred to two females from Batavia (now Jakarta). 

HampsonÕs type specimen (Hampson 1903), labelled Darjiling, is in the 

British Museum (Natural History) (BMNH); and the distribution of his 

species was given as Sikkim; Celebes, Palos B., Dongola (Doherty). 

Dongola is currently spelt Donggala and is at the southern headland of Palos 

bay at the head of which is Palu. DohertyÕs obituary (Hartent 1901) reveals 

that he collected there in August and September 1896. 

Southeast Asia China Pacific 

3 ++ Phil 2 ++ Yunnan Province 2 ++ PNG 

+ Thai 

P Brun, Indo 

The origin of mango ( Mangifera indica) 

is believed to be in the India- 

Myanmar region, from which it might be inferred that D. sublimbalis also 

evolved within this region, unless it has transferred to M. indica from a 

related plant. 

India, Myanmar, Thailand, China (Yunnan Province: Li et al. 1997), Brunei, 

Philippines, Indonesia (Java, Sulawesi, Irian Jaya), Papua New Guinea, 

Torres Strait (Dauan Is, Saibai Is: AQIS 1991; NAQS 1993) (Fenner 1987; 

Singh 1987). 

It is apparently not known in Pakistan (M.A. Poswal pers. comm. 1997), 

Sri Lanka (J. Edirisinghe pers. comm. 1997), Nepal (Neupane 1995) or 

Peninsular Malaysia (Yunus and Ho 1980; Tan Chai-Lin pers. comm. 1997) 

and does not occur on the Australian mainland or in the oceanic Pacific. It is

Biology 

4.6 

Deonalis sublimbalis 

107 

widely distributed throughout Papua New Guinea coastal mainland and 

islands (F. Dori pers. comm. 1997). 

Specimens in the BMNH carry the following labels 

India: Darjiling (now Darjeeling) 

Calcutta 22 March, 1945 

Orissa March, 1952 

Myanmar: Rangoon March, 1923 

Thailand and Philippines: no dates 

Brunei: 12 April, 1973, 5 September 1992 

Indonesia: Java August, 1922 (Koepoedan) Sulawesi 

(Minahassa, Tomohon). 

Irian Jaya July, 1936 (Cyclops Mts, 

Sabron 2000ft) 

Papua New Guinea: Kokoda August, 1933; 

and in the Australian National Insect Collection the following labels: 

Papua New Guinea: Finisterre Range 23 JuneÐ21 July, 1958 

(Gabumi, 2000ft) 

Telefomin 2 May and 18 June, 1959 

(Feramin 4700ft) 2 MayÐ18 June 

Port Moresby 5 MarchÐ12 May, 1963 

(Mt Lawes 1300ft) 

Referring presumably to the major (summer) crop of mangos, Fenner (1987) 

and Golez (1991a) comment that the eggs are oval, waxy white and 

generally laid in masses near the apex of the developing fruit. However 

F. Dori (pers. comm. 1997) reports that, in the winter (July) crop near Port 

Moresby (PNG), eggs were white to crimson. They were laid in groups of 1 

to 4 near or on the peduncle at the base of the fruit and sometimes covered by 

the sepals or deposited in small crevices in the fruit. On hatching, larvae 

travel to the apex to enter the fruit. Oviposition occurs as early as 45 to 55 

days after flower induction and continues up to fruit maturity. After an egg 

incubation period of 3 to 4 days, larvae hatch and pass through 5 instars in 

the next 14 to 20 days. The larva has a brown or black head and white body 

with red segmental bands. It feeds first in the pulp of the fruit (1st and 2nd 

instars) and later in the seeds. The tunnels formed gradually broaden as the 

larvae grow to about 2 cm in length. Fruit in all stages of development from 

marble size upwards are attacked. As many as 11 larvae may be found in a 

fruit, although there is commonly only one. A pre-pupal stage lasts about 2 

to 3 days and pupation occurs in a silk-lined earthern cocoon. In wooden


Host plants 

cages in Papua New Guinea, larvae pupated in strongly spun cocoons 

covered with particles of chewed wood, suggesting that pupation on bark 

may occur in the field. In India, Sengupta and Behura (1955, 1957) record 

pupation generally inside the fruit, the moth emerging through an exit hole. 

The pupal period lasts from 9 to 14 days, so that the total period from egg to 

adult takes from 28 to 41 days. Adult longevity is 8 to 9 days. Adult males 

can be distinguished from females by having expanded dark brown, hairy, 

mesothoracic tibiae (Leefmans and Van der Vecht 1930; Vožte 

1936; 

Kalshoven 1981; Fenner 1987, 1997; Golez 1991a). 

Adults are generally nocturnal and, during the day, spend most of their 

time resting under leaves on the tree. They are seldom attracted to light. 

Females prefer to oviposit on fruit protected from direct light. Newly 

hatched larvae stay together and tunnel into the fruit near where the eggs 

were laid. If later instar larvae are crowded, some may leave by suspending 

themselves on silken threads, which also facilitate transfer to other fruits. A 

shorter developmental period was observed for both males and females 

reared on the pulp than on the seed of carabao mangoes, although those 

reared on the seeds were larger and lived longer, females producing more 

eggs. Development differed slightly on different mango varieties (Golez 

1991a). 

In cages in Papua New Guinea only a small proportion of pupae yielded 

adults in the several months after pupation, suggesting a pupal diapause 

which may synchronise the life cycle with the seasonal fruiting of its host 

(Fenner 1987, 1997). 

The commonest host, wherever D. sublimbalis occurs, is Mangifera indica, 

but there are records also from M. odorata in Papua New Guinea and 

Indonesia from M. minor in Papua New Guinea (F. Dori pers. comm. 1997). 

and from Bouea burmanica in Thailand (Beller and Bhenchitr 1936). All 

four belong to the family Anacardiaceae (Sengupta and Behura 1955; 

Kalshoven 1981; M. Schaffer pers. comm. 1997). Larvae are unable to 

develop on parts of the mango tree other than the fruit, or in the fruit of 

avocado, chico, guava, jackfruit, papaya, santol, sineguelas or star apple 

(Golez 1991a). However, as the genus Mangifera contains many species it is 

quite possible that further wild hosts will be found (Fenner 1997). Indeed the 

label data, quoted earlier, on specimens collected well outside the major 

fruiting season of M. indica suggests that this may well be so. The genus 

Mangifera contains about 62 species of tall evergreen trees which are native 

to the area stretching from India to Papua New Guinea, with the greatest

Damage 

4.6 


109 

number in the Malay Peninsula. Fifteen species bear edible fruit, but only 

M. indica is widely planted of the 6 species sometimes cultivated. M. indica 

probably originated in the Indo-Myanmar region and grows wild in the 

forests of India, particularly in hilly areas in the northeast. It has been grown 

throughout the Indian sub-continent for at least 4000 years. It was probably 

taken to Malaysia and eastwards further into Southeast Asia between 300 

and 400 AD and there are now many commercial varieties (Purseglove 

1968). 

Mango fruit in all stages of development are attacked, often leading to 

premature drop. First and second instar larvae feed on the tissues beneath the 

skin, making tunnels towards the seed. Larger larvae destroy the seed. Soon 

after boring starts, secondary infestations of bacteria, fungi, fruit flies (e.g. 

Bactrocera ferrugineus, 

B. frauenfeldi), 

and other pests occur. Liquid 

exudes from the skin of attacked fruit at the opening of the entry tunnel and 

trickles down to the drip point where it accumulates. It rapidly darkens to 

form a characteristic black spot, often about 1cm in diameter at the tip of the 

fruit (Fenner 1987). Another common sign of borer damage is the bursting of 

the apex and longitudinal cracking of the fruit. In Guimaras Province 

(Philippines), up to 12.5% fruit infestation was recorded by Golez (1991a), 

with up to 14.5 larvae occurring per kg fruit. In years of serious infestation, 

yield could be reduced by as much as 40 to 50 per cent (Tipon 1979). In 

Papua New Guinea, fruit infestation levels of greater than 20% are 

encountered in the Port Moresby area (F. Dori pers. comm. 1997; T.L. 

Fenner pers. comm. 1997). In India the seeds are used as human food in 

times of famine and a flour is made from them (Purseglove 1968). 

However, for an insect that can be significantly damaging to mango fruit, 

it is remarkable that there are so very few references to it in the literature. It 

is, perhaps, instructive to list those that are directly relevant, so as to 

contribute to determining (i) whether it has, for many years, frequently been 

overlooked as a pest, (ii) whether it has spread to new areas in recent times, 

(iii) whether it has only become a pest of edible mangoes in recent times, (iv) 

whether suppression by natural enemies is no longer as effective as it once 

was and/or (v) whether there are other reasons. 

D. sublimbalis must have been present in northern India before it was 

described in 1903 (Hampson 1903), although it was not mentioned in the 

books by Maxwell-Lefroy and Howlett (1909), Fletcher (1914) or Ayyar 

(1963) all dealing with insects of agricultural importance in India. It was, 

however, reported a little later by Wadhi and Batra (1964) who referred to


papers by Sengupta and Behura (1955, 1957) and Sengupta and Misra 

(1956). It was also reported, briefly, by Nair (1975) and, in more detail, by 

Butani (1979). Strangely, the above Sengupta and Behura (1955) reference 

lists D. albizonalis among new records of crop pests in Orissa and then only 

of grafted mangoes in Puri District, implying that it was not known much 

earlier there as a damaging species. Furthermore, only recently 

(Zaheruddeen and Sujatha 1993) was D. sublimbalis recorded as having 

caused serious losses to mango fruits from marble size to maturity in 

Godavari Districts of Andhra Pradesh. 

Although a specimen was collected in Rangoon in 1923 (BMNH), 

D. sublimbalis was not listed by Ghosh (1940) in his major work ÔInsect 

Pests of BurmaÕ or by Yunus and Ho (1980) in Malaysia when dealing with 

economic pests from 1920 to 1978. This striking absence of records from 

peninsular Malaysia continues to this day (Tan Chai-lin pers. comm. 1997). 

Nevertheless, D. sublimbalis has been well known in Thailand since 1936 

(Beller and Bhenchitr 1936; Cantelo and Pholboon 1965; Wongsiri 1991; 

Kuroko and Lewvanich 1993). 

In the Philippines it was not recorded by Cendana et al. (1984) in ÔInsect 

Pests of Fruit Plants in the PhilippinesÕ, so it was evidently not generally 

regarded as a pest at that time, although a paper recording 40 to 50% damage 

in bad years had been delivered 5 years earlier (Tipon 1979). A 

comprehensive account of up to 12.5% infestation of fruit in Guimaras 

Province was published in 1991 by Golez (1991a,b). 

In contrast, in Indonesia it was present prior to 1899 (Snellen 1899) and 

has been well known as a mango pest since 1930 (Leefmans and van der 

Vecht 1930). Its damaging presence there is also documented by Vote 

(1936) and Kalshoven (1981). 

D. sublimbalis has been known in Irian Jaya since 1936 (BMNH 

specimen) and was common in mangoes in Jayapura in the early nineties 

(T.L. Fenner pers. comm. 1997). It was collected in Papua New Guinea 

(Kokoda) in 1933 (BMNH specimen) and was recorded again in 1958, 1959 

and 1963 (ANIC specimens) and is common nowadays in Port Moresby. It 

was first recorded on Australian islands in Torres Strait (Saibai I) in 1990 

and again in October 1996 (at a level of about 1% infestation on Dauan I) 

(Australian Quarantine Inspection Service).


Table 4.6.1 

4.6 


111 

Leefmans and van der Vecht (1930) commented that no parasites had been 

bred in their studies on D. albizonalis in Java. 

In Luzon (Philippines) the egg parasitoids Trichogramma chilonis and 

T. chilotraeae (Table 4.6.1) were recorded by Golez (1991a) who reported, 

however, that no parasitoids were encountered at that time in the three 

municipalities of Guimaras, all of which had dry, dusty and windy 

conditions. 

Golez (1991a) reported that predation in Guimaras occurs as larvae leave 

the fruit, either to migrate to another fruit or to pupate in the soil. The most 

important predator was the vespid Rhychium attrisium which appeared to be 

the main cause of the high larval disappearance that occurs. R. attrisium is 

abundant in summer, especially during warm sunny days. 

Larvae were attacked by a fungus in the laboratory in Indonesia 

(Leefmans and van der Vecht 1930). The wasp Evania appendigaster is 

reported as a larval/pupal parasite (Golez 1991b), but Fenner (1997) points 

out that this record needs confirmation since the Evaniidae are reportedly all 

parasites of cockroach eggs. 

A tachinid, Carcelia ( Senometopia) 

sp., was reared from mango fruit 

possibly infested with D. sublimbalis near Nodup in September 1982 

(J. Ismay pers. comm. 1997) and also from a D. sublimbalis larva near 

Rabaul (both Papua New Guinea) in 1984 (F. Dori pers. comm. 1997). 

Natural enemies of Deanolis sublimbalis 

Species 

DIPTERA 

TACHINIDAE 

Location Reference 

Carcelia sp. 

HYMENOPTERA 


Rabaul (PNG) F. Dori pers. comm. 1997 

Trichogramma chilonis 

Philippines Golez 1991a 

Trichogramma chilotraeae 

EVANIIDAE 

Philippines Golez 1991a 

Evania appendigaster 

VESPIDAE 

Golez 1991b 

Rhychium attrisium 

Philippines Golez 1991a


Comment 

It is tempting to postulate that the damage that is actually due to the red 

banded mango caterpillar is commonly attributed to other causes. Perhaps 

this is due, in part, to the fact that larvae have often left the fruit before the 

cause of damage is investigated although, with a larval duration of 2 to 3 

weeks, this would be surprising, particularly when there is a characteristic 

dark spot for much of this time at the drip point of the mango fruit. 

The absence of records of its presence over vast areas within its 

distribution range suggests that its abundance must be very low (perhaps due 

to inhospitable host varieties or effective biological control) or, perhaps, that 

it does not occur there. 

The only record of effective chemical control is of 4 applications of 

cyfluthrin or deltamethrin at 60, 75, 90 and 105 days after fruit induction 

(Golez 1991a). 

Further research for natural enemies attacking eggs, larvae and pupae 

within its long established range would be necessary to determine whether 

any are likely to be promising for biological control. 

If, as is very probable, D. sublimbalis produces a sex pheromone, its 

availability as a lure would be of great value as a means of monitoring the 

presence and distribution of the red banded mango caterpillar in mango and 

other hosts. Its identification, synthesis and availability should have high 

priority.

4.7 Diaphorina citri 

India 

20° 

Myanmar 

P Laos 

0° 

20° 

China 

++ 

Thailand 

P 

Cambodia 

Vietnam 

++ 

+ Brunei 

Malaysia 

+ 

Singapore 

++ 

Indonesia 

Taiwan 

+++ 

++ 

Philippines 

Australia 

+ 

Papua 

New Guinea 

113 

The citrus psyllid Diaphorina citri is native to the Indo-Malaysian region, but has spread 

outside it to RŽunion, Mauritius, Saudi Arabia, Honduras, and Brazil. The sap it removes 

from new flushes of citrus growth is of minor consequence, but it is the vector of a 

devastating bacterial disease, citrus greening. 

Its major controlling factors are high rainfall (washing off eggs and young nymphs) and 

two parasitoids, Tamarixia radiata and Diaphorencyrtus aligarhensis. 

Where these 

parasitoids are native they are very heavily attacked by hyperparasitoids which diminish 

their effectiveness. Freed of these hyperparasitoids T. radiata has been established in 3 

countries where it was not present: RŽunion, Madagascar and Taiwan, resulting in 

excellent biological control. Although T. radiata appears to be widespread in Southeast 

Asia, observations might well disclose regions where it is not present and could be 

introduced with advantage. The prospects for successful biological control of D. citri are 

good when it invades regions where hyperparasitoids of T. radiata are absent or deficient. 

20° 

0° 

20°


Diaphorina citri Kuwayama 

Rating 

Origin 

Distribution 

Hemiptera, Psyllidae 

citrus psyllid, Asian citrus psyllid 


8 ++ Viet, Indo, Phil 3+++ absent 

+ Msia, Sing 

P Myan, Thai 

These Southeast Asian ratings arose from an earlier survey of country 

opinions (Waterhouse 1993b) and may not reflect current assessments. 

The Indo-Malaysian region. D. citri was described from Punjab, India 

(Waterston 1922). There is evidence of recent spread into the southeastern 

and eastern portions of Southeast Asia. 

D. citri is widespread from Afganistan eastwards through Pakistan, India, 

Nepal and Bhutan to Southeast Asia, southern China (up to about 30¡N, Xie 

et al. 1988), Taiwan (Catling 1970; Tsai et al. 1984; Aubert 1990) and the 

Ryuku Is (Japan) (Miyakawa and Tsuno 1989). It has recently become 

established in Ende (Flores) and Timor and in Irian Jaya (Aubert 1989b, 

1990). D. citri was collected in June 1993 in the Jayapura area of Irian Jaya 

and citrus there showed symptoms of greening (Northern Australia 

Quarantine Strategy 1993). It has been introduced to RŽunion, Mauritius, 

Comoro Is (Hollis 1987), Saudi Arabia (Wooler et al. 1974), Yemen (BovŽ 

1986), Brazil (Silva et al. 1968; Bergmann et al. 1994) and Honduras 

(Burckhardt and Martinez 1989). In 1990 there were still limited areas free 

of D. citri in east Mindoro (Philippines) and Palau and Tambun (Malaysia). 

It is not yet recorded from Papua New Guinea and does not occur in 

Australia, the Oceanic Pacific or North America. 

In RŽunion it has not colonised citrus plantings above 800 m, where the 

lowest temperature is 7¡C, whereas in Malaysia the height limit is 1200 m 

with a minimum temperature of 14¡C.

Biology 

Table 4.7.1 

4.7 


115 

D. citri survives a wide range of temperature extremes from 45¡C in Saudi 

Arabia to Ð7¡ to Ð8¡C in China, thereby tolerating cold that will kill citrus 

(Xie et al. 1989a). Far more than temperature, high humidity and rainfall are 

important limiting factors, rain by washing off eggs and early instar nymphs 

and humidity by favouring fungal attack. These two factors are mainly 

responsible for the low D. citri populations on the windward (rainy) side of 

Mindoro (Philippines) and RŽunion (Aubert 1989a). 

There have been several studies on the life cycle of D. citri, 

which 

conform generally with the results in Table 4.7.1, leading to up to 11 

generations a year in Fujian Province, China (Xu et al. 1988b, 1994). D. citri 

has a short life cycle and high fecundity and is commonest in hot coastal 

areas. Mating commences soon after the insects become adult and, after a 

pre-oviposition period of about 12 days, eggs are laid singly inside halffolded 

leaves of buds, in leaf axils and other places on the young tender 

shoots. Average adult lifespan is 30 to 40 days, although overwintering 

adults had a lifespan of 260 days (Xu et al. 1994). 

Bionomics of D. citri (average in days) in Fujian Province, 

China (Xu et al. 1988b) 

Adult life-span 

Max Min 

Eggs per 

female 

Incubation Nymphal 

development 

EggÐadult 

Spring 96 28.1 17.7 10 31.8 42 

Summer 46 19.7 43.8 2 10.3 13 

Autumn 59 31.6 22.6 4 16.8 21 

Winter 131Ð165 

The abundance of both eggs and nymphs is correlated with the availability of 

new growth flushes and breeding is largely suspended when trees become 

dormant. On its favoured host plant Murraya paniculata in Fujian, 

populations may average 51 adults per young shoot and a 4-year-old plant 

produces 900Ð1000 shoots. On mandarin ( Citrus reticulata) 

the average 

colony size is 20 per shoot, with 600 to 650 shoots, and peak abundance 

occurs about 6 weeks later than on M. paniculata (Aubert 1990). D. citri 

nymphs develop well under cool, humid spring conditions, but are seriously 

affected by fungal infections under hot, humid conditions. On 

M. paniculata, 

adult numbers were highest on leaf midveins (43%), 

followed by petioles (30.7%), leaf blades (23.7%) and stems (2.6%) (Tsai et 

al. 1984).


Host plants 

Damage 

D. citri nymphs normally lead a sedentary existence clustered in groups, 

but will move away when disturbed. Adults are 2.5 mm long and jump when 

disturbed, whereupon they may fly up to 5 m before settling again. Seasonal 

migratory flights occur when adults fly up to about 7 m above ground level, 

entering mild winds which may carry them up to 4 km distant (Aubert 1990). 

Flying adults are attracted to yellow traps, which have been used for 

sampling (Aubert and Xie 1990). Adult D. citri have yellowish-brown 

bodies, greyish-brown legs and transparent wings. They have white spots or 

are light brown with a broad, beige, longitudinal, central band. 

D. citri feeds and breeds on the entire group of horticultural Citrus, 

with 

additional hosts in eight different genera belonging to the Aurantoidea 

(Aubert 1990). D. citri thus has a wider host range than the greening 

organism it transmits to citrus (see Damage). An indication of the relative 

suitability of its various host plants is shown in Table 4.7.2, although there 

may be local modifications of the groupings. This is probably due to 

different D. citri biotypes. For example, unlike RŽunion populations, 

Malaysian populations breed well on Bergera koenigii and, in the 

Philippines, adults are more attracted by Clausena anisumolens than by 

Murraya paniculata (Aubert 1990). Overall, jasmin orange, Murraya 

paniculata, is the preferred host and this plant is widely grown in Southern 

and Southeast Asia as an ornamental shrub and hedge plant. 

Although sap removal by large populations of D. citri can cause young 

foliage on flushes of growth to wilt, by far the most damaging effect of 

feeding is due to the transmission of a gram-negative bacterium which is the 

cause of citrus greening, known as huanglungbin in China (Xu et al. 1988a). 

Citrus greening is known to affect 3 genera of the subtribe Citrinae, namely 

Citrus, 

Poncirus and Fortunella (Aubert 1990). It has also been 

experimentally transferred from Citrus to Madagascar periwinkle 

( Catharanthus roseus (Ke 1987). Once infected with the bacterium, D. citri 

remains infective for its lifetime, but does not pass on the infection 

transovarially. Amongst citrus, pummelo and lemon are less affected by 

greening than most other species. D. citri is the only known vector of citrus 

greening in Asia, although several other psyllids attacking citrus have been 

described: D. auberti (Comoro Is: Hollis 1987), Psylla citricola, 

P. citrisuga and Trioza citroimpura (China: Yang and Li 1984) and Psylla 

murrayii (Malaysia: Osman and Lim 1990).

Table 4.7.2 

Preferred host plant 

Good host plants 

Common host plants 

Occasional host plants 

Diaphorina citri host plants (after Aubert 1990) 

Murraya paniculata ( jasmin orange) 

Citrus aurantifolia (lime) 

Bergera (Murraya) koenigii (curry bush) 

Leaf sucking Egg laying Nymphal 

development 

+++ +++ +++ 

+++ +++ +++ 

Citrus limon (lemon) 

Citrus sinensis (sweet orange) 

Citrus medica (citron) 

Citrus reticulata (mandarin) 

Microcitrus australisiaca* 

Citrus maxima var. racemosa (pummelo) 

Citrus hystrix ( Mauritius papeda) 

Citrus madurensis 

Clausena excavata 

Clausena lansium 

++ ++ ++ 

Citrus maxima (pummelo) + + + 

Triphasia trifoliata* 

+ + + 

Fortunella sp.* (kumquat) + + + 

Poncirus trifoliata* 

+ + Ð 

Clausena anisumolens (anise) + + + 

Merrillia caloxylon* 

+ Ð Ð 

Toddalia asiatica* 

+ Ð Ð 

4.7 


117

Table 4.7.2 (contÕd) Diaphorina citri host plants (after Aubert 1990) 

Occasional host plants 

Leaf sucking Egg laying Nymphal 

development 

Vepris lanceolata* 

+ Ð Ð 

Swinglea glutinesa* 

+ unknown unknown 

Atalantia sp. + unknown unknown 

Clausena indica* 


Murraya exotica* 


Citrus species hosts are, according to the classification of Jones (1990): 

+++ very common; 

++ usual 

+ occasional; 

Ð complete life cycle not observed 

*observations on caged insects 


4.7 Diaphorina citri 119 

In Africa, RŽunion, Madagascar and Saudi Arabia another psyllid Trioza 

erytreae transmits a slightly different citrus greening organism (see later 

under RŽunion). 

Citrus greening is believed to have originated in northeastern 

Guangdong Province (Lin and Lin 1990). Amongst other symptoms, the 

leaves of new green shoots first turn yellow at their base, then often become 

mottled yellow and drop. The branches remain small, upright and stiff. 

Diseased trees flower abundantly in the off-season and flowers drop readily 

or result in small, irregular fruit whose base turns red before the remainder 

changes from green (Ke 1987). Citrus greening is widespread throughout 

South and Southeast Asia, where it is almost always the most serious disease 

of citrus. It is spread to new areas by infected nursery plants or infected 

budwood and within orchards by D. citri (Capoor et al. 1967; Whittle 1992). 

However, D. citri has been intercepted by quarantine in France on citrus 

imported from Honduras (Burckhardt and Martinez 1989). The tonnage of 

citrus produced worldwide is second as a fruit crop only to that of grapes 

(Aubert 1987b). An extremely serious citrus disease which already affects 

nearly 50 countries in Asia and Africa must, therefore, be regarded as of 

major importance. It is reported that a total of over a million trees are 

destroyed each year in China, Thailand, Malaysia, Indonesia and Philippines 

alone (Aubert 1987a). In Indonesia citrus greening has caused the loss of 

many millions of trees. Small farmers are frequently reluctant to remove 

declining trees before they almost cease bearing. This tends to increase 

D. citri populations, which breed on young flush since a symptom of 

greening is unseasonal flushing (Whittle 1992). The recent history of 

production in northern Vietnam, where citrus is grown mainly in larger 

orchards or state farms, is typically cyclical, with the gradual destruction of 

trees by greening and then wholesale removal and replanting. A new cycle of 

planting commenced in the late 1980s, but greening is already to be seen in 

many young orchards, although populations of D. citri are still low (Whittle 

1992). Only by keeping populations at very low levels by biological control 

and/or insecticides will the rate of spread of greening be diminished. 

Insecticides are said to be highly cost effective if used only during a 

restricted flushing period, but if needed frequently they are very costly and 

environmentally undesirable. Recent developments with carefully specified, 

highly refined petroleum oils has given high levels of control of D. citri 

(A. Beattie pers. comm. 1995), with presumably little direct effect on its 

parasitoids. Whittle (1992) reported that he was unable to find D. citri in the 

vicinity of Ho Chi Minh City (southern Vietnam), a very unusual situation 

for an area with a fairly long history of citrus cultivation.


The Asian citrus greening bacterium can withstand high temperatures 

and occurs in China, Southeast Asia, India and Saudi Arabia. On the other 

hand, Southern African greening, which is transmitted by the psyllid Trioza 

erytreae, is heat-sensitive and symptoms do not develop in climates where 

temperatures above 30¡C are recorded for several hours a day. In addition to 

Southern Africa, this greening occurs also in North Yemen (Garnier et al. 

1988). 


Identified natural enemies are listed in Table 4.7.3. There are also reports of 

a number of unidentified predators (coccinellids, chrysopids, mantids, 

spiders). It is noteworthy that only 2 primary parasitoidsÑboth attacking 

D. citri nymphsÑhave so far been recorded, the widespread endoparasitic 

encyrtid Diaphorencyrtus aligarhensis and the more restricted ectoparasitic 

eulophid Tamarixia radiata, which has been introduced to several countries 

for biological control. Both feed on the haemolymph of some hosts, resulting 

in their death, as well as using other hosts for oviposition. 

Where they occur naturally, both D. aligarhensis and T. radiata are 

heavily attacked by a wide range of hyperparasitoids (Table 4.7.4). Of these, 

Tetrastichus sp. is the most important for T. radiata, causing an average of 

21.8% parasitisation in 1988 (rising to a maximum of 87.9%) and 28.7% in 

1989 in Fujian Province, China. Chartocerus walkeri (9.3% in 1988 and 

13.2% in 1989) is the most important for D. aligarhensis (Table 4.7.5). 

A valuable illustrated guide to the hyperparasitoids associated with 

D. citri is provided by Qing and Aubert (1990).

Table 4.7.3 Natural enemies of Diaphorina citri (* indicates introduced to this country) 

Species Region Reference 

HYMENOPTERA 

ENCYRTIDAE 

Diaphorencyrtus aligarhensis 

(= Aphidencyrtus diaphorinae 

= Diaphorencyrtus diaphorinae 

= Psyllaephagus diaphorinae 

= Aphidencyrtus aligarhensis) 

EULOPHIDAE 

Tamarixia radiata 

(= Tetrastichus radiatus) 

India 

Vietnam 

Taiwan 

Shafee et al. 1975; Hayat 1981 

Myartseva & Tryapitzyn 1978; 

van Lam 1996 

Prinsloo 1985 

Lin & Tao 1979 

Comores Is Aubert 1984b 

RŽunion Aubert & Quilici 1984, Quilici 1989 

Philippines Prinsloo 1985; Gavarra & Mercado 1989; 

Gavarra et al. 1990 

China Tang 1989 

Indonesia 

Malaysia 

India 

RŽunion* 

Nurhadi 1989; Nurhadi & Crih 1987 

Lim et al. 1990 

Waterston 1922; Husain & Nath 1924; Quilici 1989, 

Etienne and Aubert 1980 

Saudi Arabia Aubert 1984a 

Mauritius* Aubert 1984c 

Nepal Lama et al. 1988; Otake 1990 

Taiwan* Chiu et al. 1988 

China Liu 1989; Tang 1989; Qing & Aubert 1990 

Indonesia 

Malaysia 

Nurhadi & Crih 1987; Nurhadi 1989 

Lim et al. 1990 

Thailand Qing & Aubert 1990 

Vietnam Myartseva & Trijapitzyin 1978; 

van Lam 1996 

4.7 Diaphorina citri 121

Table 4.7.3 (contÕd) Natural enemies of Diaphorina citri (* indicates introduced to this country) 

Species 

COLEOPTERA 

Region Reference 

COCCINELLIDAE 

Cheilomenes sexmaculata China Xia et al. 1987 

NEUROPTERA 

CHRYSOPIDAE 

Chrysopa boninensis China Liu 1989 

ARACHNIDA 

SALTICIDAE 

Marpissa tigrina India Sanda 1991 

FUNGI 

Beauveria bassiana China Chen et al. 1990 

Cephalosporium (= Verticillium) lecanii China Xie et al. 1988 

Fusarium lateritium China Xie et al. 1988 

Paecilomyces sp. China Xie et al. 1988 


Table 4.7.4 Hyperparasitoids of Diaphorina citri (mostly after Tang 1989) 

Hyperparasitoid 

EULOPHIDAE 

Attacks Region Reference 

Tetrastichus sp. T.r. & D.a. China Tang 1989 

D.a. Taiwan Hayat & Lin 1988; Chien et al. 1989 

ENCYRTIDAE 

Philippines Balthazar 1966, unpublished 

Syrphophagus taiwanus T.r. & D.a. Taiwan Hayat & Lin 1988; Chien et al. 1989 

T.r. & D.a. China Tang 1989 

Ageniaspis sp. D.a. Taiwan Hayat & Lin 1988; Chien et al. 1989 

Cheiloneurus sp. 

D.a. 

Taiwan Hayat & Lin 1988; Chien et al. 1989, 

? 

Philippines Baltazar 1966, unpublished 

?Psyllaephagus sp. 

T.r. & D.a. China 

Tang 1989 

Philippines Balthazar 1966, unpublished 

Tang 1989 

Several unidentified 

SIGNIPHORIDAE 

D.a. China Tang 1989 

Chartocerus walkeri T.r. & D.a. Taiwan Hayat & Lin 1988; Chien et al. 1989 

T.r. & D.a. China 

Tang 1989 

Signiphora sp. 

PTEROMALIDAE 

D.a. Gavarra et al. 1990 

Pachyneuron concolor 

APHELINIDAE 

T.r. & D.a. Taiwan Hayat & Lin 1988; Chien et al. 1989 

Coccophagus ceroplastae D.a. Taiwan Hayat & Lin 1988; Chien et al. 1989 

Coccophagus sp. D.a. Taiwan Hayat & Lin 1988; Chien et al. 1989 


Table 4.7.4 (contÕd) Hyperparasitoids of Diaphorina citri (mostly after Tang 1989) 

Hyperparasitoid Attacks Region Reference 


Marietta leopardina 

(= Marietta javensis) 

T.r. = Tamarixia radiata D.a. = Diaphorencyrtus aligarhensis 

T.r. & D.a. 

D.a. 

Taiwan 

Philippines 

Encarsia spp. T.r. & D.a. Taiwan 

China 

Unidentified sp. T.r. & D.a. Taiwan Chien et al. 1989 

Hayat & Lin 1988; Chien et al. 1989 

Balthazar 1966, unpublished 

Hayat & Lin 1988; Chien et al. 1989 

Tang 1989 


Table 4.7.5 Hyperparsitoids of Tamarixia radiata and Diaphorencyrtus aligarhensis in Fujian and Taiwan (after Qing 

1990) 

T. radiata 

Percentage of hyperparasitisation 

D. aligarhensis 

Hyperparasitoid 

EULOPHIDAE 

Fujian Fujian Taiwan Fujian Fujian Taiwan 

Tetrastichus sp. 

PTEROMALIDAE 

21.82 28.65 0.01 2.90 3.68 

Pachyneuron concolor 

SIGNIPHORIDAE 

0.45 18.50 

Chartocerus walkeri 

ENCYRTIDAE 

0.08 1.09 0.03 9.26 13.16 13.50 

Syrphophagus taiwanus 0.05 1.09 4.21 6.80 

?Psyllaephagus sp. 0.04 0.10 10.35 6.58 

Cheiloneurus sp. 

Ageniaspis sp. 

0.01 

unidentified sp.A 

Sp.B 

Sp.C 

Sp.D 

APHELINIDAE 

Encarsia sp. near transvena 

(= E. shafeei) 

3.45 

0.91 

0.18 

0.18 

0.26 

0.11 0.80 

Encarsia sp. A 0.08 0.10 0.91 1.05 

Encarsia sp. B 0.22 0.20 0.91 


Table 4.7.5 (contÕd) Hyperparsitoids of Tamarixia radiata and Diaphorencyrtus aligarhensis in Fujian and Taiwan (after Qing 

1990) 

Percentage of hyperparasitisation 

T. radiata D. aligarhensis 

Hyperparasitoid Fujian Fujian Taiwan Fujian Fujian Taiwan 


Marietta leopardina 0.25 2.50 

Coccophagus ceroplastae 0.01 

Coccophagus sp. 0.10 

Unidentified sp. 0.05 0.01 

Totals 22.24 30.14 0.90 30.14 28.94 39.72 



It is noteworthy that T. radiata, which has fairly recently (1984Ð1988) 

been introduced into Taiwan, was hyperparasitised to the extent only of 

0.95% in 1989, whereas 42.2% of the native D. aligarhensis was attacked 

(Qing 1990). 

The levels of hyperparasitisation of both primary parasitoids seriously 

affects their capacity to develop high populations and hence to produce 

maximum reduction of host populations. Nevertheless, each primary 

parasitoid killed is also a D. citri killed, so the overall mortality of D. citri is 

the sum of the mortalities produced by both primary parasitoids and their 

hyperparasitoids. It is abundantly clear that all hyperparasitoids must be 

rigorously excluded when transferring primary parasitoids from one region 

to another. 


CHINA 

The parasitoid Tamarixia radiata, obtained originally from India, has been 

used in successful biological control projects in RŽunion, Mauritius and 

Taiwan and in an attempt in the Philippines (Table 4.7.6). These projects and 

comments on the situation in several other countries follow. 

Table 4.7.6 Introductions for the biological control of Diaphorina citri 

Species 

EULOPHIDAE 


Tamarixia radiata India RŽunion 1978 + Aubert & Quilici 1984; 

Quilici 1989 

RŽunion Mauritius after 1978 + Quilici 1989 

RŽunion Taiwan 1983Ð86 + Chiu et al. 1988; 

Chien et al. 1988 

RŽunion Philippines 1989 + 

? 

Gavarra et al. 1990 

Mercado et al. 1991 

In Guangdong, predators (lacewings, ladybird beetles, thrips, spiders) 

caused about 80% mortality of D. citri. Duration of daylight (short days 

reducing oviposition), quality of the flushes, and pesticide usage were other 

important factors influencing D. citri populations (Chen 1988). It appears 

that some Chinese farmers may spray citrus up to 50 times a year. 

In Fujian there are 8 generations a year of D. citri on jasmin orange, 

Murraya paniculata and populations reach their peak in summer and early 

autumn during hot, dry weather when fresh shoots appear regularly. 

Populations are lowest in cold, wet weather with average temperatures of


9.1¡ to 12.2¡C. Rainfall affects populations since eggs are laid on very young 

twigs and are easily washed off. A Tamarixia sp. was recorded in September 

1987 and caused 83.3% parasitisation of nymphs in late autumn. In spring 

1988 its population was low, but D. citri mainly overwinters as the adult and 

Tamarixia only attacks nymphs. Predators included coccinellids (especially 

Cheilomenes sexmaculata and Harmonia axyridis), lacewings, spiders and 

praying mantids (Xia et al. 1987; Ke 1991). 

In Guangdong a maximum of 75% mortality of D. citri was recorded as 

being due to the hyperparasitoid Tetrastichus sp. (Liu 1989). 

Beauveria bassiana (Chen et al. 1990a), Cephalosporium (Verticillium) 

lecanii and two other fungi (Fusarium lateritium and Paecilomyces sp.) 

were found attacking D. citri. Suspensions of C. lecanii sprayed on to 

D. citri displayed a very high pathogenicity (Xie et al. 1988, 1989b). 

INDONESIA 

Citrus greening is also known as citrus vein phloem degeneration. In East 

Java, both T. radiata (the commoner) and D. aligarhensis (the more 

widespread) were found in 1987 attacking D. citri on Murraya paniculata 

(Nurhadi and Crih 1987). D. citri is known to occur in Irian Jaya and may 

have been introduced in recent times, but it is not known if it is parasitised 

there. 

MALAYSIA 

Both Tamarixia radiata and Diaphorencyrtus aligarhensis are present with 

parasitisation rates ranging up to 28% in 4th and 5th instar nymphs (Osman 

and Quilici 1991) or up to 36% parasitisation (Lim et al. 1990). T. radiata is 

also present in Sarawak (S. Leong, pers. comm. 1995). 

NEPAL AND BHUTAN 

Both T. radiata and D. aligarhensis are present in some parts of both 

counties and may cause parasitisation of D. citri in excess of 90% (Lama and 

Amtya 1991; Lama et al. 1987). 

PHILIPPINES 

Citrus greening, also known as citrus leaf mottle, was already causing 

serious damage in the early 1960s. However, as late as 1988, the windward 

side of Mindoro island with an average rainfall of 3000 mm was virtually 

free of D. citri and citrus greening, presumably due to the adverse effects of 

high rainfall (Aubert 1989a). D. aligarhensis was reared from D. citri 

(25.7% parasitisation) and also 4 hyperparasitoids (Marietta sp. and 3 

unidentified species), resulting in an overall mortality of D. citri of 48.3%. A 

Beauveria sp. attacked many psyllids and in turn was parasitised by another 

ascomycete, probably a Melanospora sp. (Gavarra and Mercado 1989). 

Later (Mercado et al. 1991), up to 62.2% parasitisation by D. aligarhensis

RƒUNION 


was reported in Mindoro. In another study, Gavarra et al. (1990) recorded 

that, in addition to the primary parasitoid D. aligarhensis (17.6 to 36.1% 

parasitisation), 5 hyperparasitoids were reared from D. citri: Marietta 

leopardina (= M. javensis), Tetrastichus sp., Psyllaephagus sp., Chiloneurus 

sp. and Signiphora sp. 

Because it was apparently absent from the Philippines (Baltazar 1966), 

Tamarixia radiata was introduced from RŽunion in 1988, but attempts to 

rear it failed. A second consignment late that year was soon followed by the 

discovery of it nearby in the field in April 1989, with recoveries continuing 

in 1990 (Gavarra et al. 1990). However, Mercado et al. (1991) expressed 

some doubts that it had become established. It is thus not clear whether 

T. radiata ever occurred naturally in the Philippines. 

The rainy, windward, east side of RŽunion has much lower D. citri 

populations and citrus trees there are much less exposed to transmission of 

greening (Aubert 1989a). Quilici (1989) has provided a valuable overview 

of the biological control of citrus psyllids in RŽunion. In the early 1970s, 

RŽunion and Mauritius were the only places known where Diaphorina citri 

and Trioza erytreae, the two psyllid vectors of citrus greening disease, 

occurred (Aubert 1987c). (Both are now known also from Saudi Arabia and 

Yemen: BovŽ 1986). Both psyllids were abundant in RŽunion and Mauritius 

and citrus greening was seriously affecting citrus production in both islands. 

The Asian citrus psyllid D. citri was most abundant below 500 m in the 

hotter and drier leeward side of RŽunion, where the average rainfall is below 

1000 m. On the other hand, the drought-sensitive African psyllid T. erytreae 

was particularly abundant in the cooler, moister regions above 600 m. The 

only nymphal parasitoid of both species was the relatively ineffective 

D. aligarhensis. Several predators exerted little control. 

Tamarixia dryi was introduced in 1974 from South Africa and, after 

elimination of hyperparasitoids, was mass produced and released in 

neglected, unsprayed citrus orchards colonised by D. citri. Populations of 

Trioza erytreae diminished progressively from 1979 to 1982, since when 

T. erytreae has not been recorded, although Tamarixia dryi is still abundant 

on another psyllid, Trioza litseae (= T. eastopi). 

In 1978 Tamarixia radiata was introduced from India and released on 

the leeward (west) side of RŽunion. From 1982 onwards D. citri has virtually 

disappeared from commercial citrus orchards, although on Murraya 

paniculata hedges there persist low populations of D. citri which are 

parasitised by T. radiata and occasionally, especially at higher altitudes, by 

D. aligarhensis.


The excellent success of these two biological control projects is ascribed 

to 3 factors: 

(i) the absence of hyperparasitoids of the primary parasitoids of Tamarixia 

dryi and T. radiata. 

(ii) the presence of an alternative host for Tamarixia dryi, which enabled it 

to maintain itself as Trioza erytreae populations diminished. 

(iii) the maintenance on Murraya paniculata hedges of low populations of 

D. citri, heavily parasitised by both T. radiata and D. aligarhensis 

(Aubert 1987c; Etienne and Aubert 1980; Quilici 1989). 

SAUDI ARABIA AND YEMEN 

In Saudi Arabia, both D. citri and Trioza erytreae are present; the former is 

the main vector of citrus greening. Both vectors are also present in Yemen 

where citrus greening at high elevations is probably the African form 

transmitted by T. erytreae (BovŽ 1986). In Saudi Arabia lime and lemon 

trees are favoured hosts of D. citri (Wooler et al. 1974). 

TAIWAN 

The nymphal ectoparasitoid Tamarixia radiata was introduced from 

RŽunion and, after mass rearing, released widely in citrus orchards and on 

Murraya paniculata hedges from 1984 to 1988. It became established, 

attaining parasitisation rates of up to 100%. Hyperparasitisation was 

initially, in 1988, below 1% (Chien 1989; Chien et al. 1988; Su and Chen 

1991), but by 1991 had risen gradually to 5.6% (Chien et al. 1991a). This is 

in contrast with levels of 72% by some 10 species attacking the native 

Diaphorencyrtus aligarhensis. High levels of attack on D. aligarhensis is 

one reason why this species is far less effective against the citrus psyllid than 

the introduced T. radiata (Chien et al. 1988). T. radiata was capable of 

maintaining D. citri at low densities in relatively stable habitats where 

Murraya paniculata was occasionally present, whereas D. aligarhensis has 

adapted to unstable habitats. However, it only provides partial control due to 

25.5 to 51.1% hyperparasitisation throughout the island. In the Taichung 

area, T. radiata was more abundant than D. aligarhensis, but the peak 

abundance of the two did not overlap and the total parasitisation varied from 

80 to 100% from February to April and 32 to 80% for the remainder of the 

year. Application of methomyl gave good control of D. citri, but it reduced 

parasitisation to a level of 0 to 4%. In an untreated citrus orchard with only 

0.1 to 0.4 D. citri adults per 10 cm length branch, the parasitoids caused 15.5 

to 46.7% parasitisation (Chien et al. 1991a). Citrus greening in Taiwan is 

known as likubin or leaf mottle disease.

VIETNAM 


Tamarixia radiata was found parasitising 3 to 10% of 4th and 5th instar 

nymphs of D. citri and Diaphorencyrtus aligarhensis was also present 

(Myartzeva and Trijapitzyin 1978; Trung 1991; van Lam 1996). 


HYMENOPTERA 

Diaphorencyrtus aligarhensis Hym.: Encyrtidae 

This primary endoparasitoid was described by Shafee et al. (1975), from 

India as Aphidencyrtus aligarhensis. Its hosts include Diaphorina citri, 

D. auberti, D. cardiae and Psylla sp. (Qing and Aubert 1990). 

The D. citri mummy parasitised by D. aligarhensis is brownish and 

hemi-spherical and encloses the parasitoid pupa. The parasitoid emerges 

from the side of the abdomen. Development from egg to adult takes 18 to 23 

days at 25 ± 1¡C and 80 to 85% relative humidity (Tang and Huang 1991). 

No males occur and unmated females produce females. On average, 4.5 eggs 

are laid per day with an average production of 144 per female. Third and 4th 

instar D. citri nymphs are preferred over 2nd instar, and 1st and 5th instars 

are not parasitised. Usually only one egg is inserted into each host, but the 

haemolymph of many young nymphs is consumed leading to their death 

(Tang and Huang 1991). 

Tamarixia radiata Hym.: Eulophidae 

This ectoparasitoid was described from India (Waterston 1922) where it is 

an important species (Husain and Nath 1924). It has been recorded in China 

(in 1982: Tang 1989), Indonesia (Nurhadi 1989), Malaysia (Lim et al. 1990), 

Nepal (Lama et al. 1988), Saudi Arabia (Aubert 1984a), Thailand (Qing and 

Aubert 1990) and Vietnam (Myartzeva and Trijapitzyin 1978). T. radiata 

has been introduced to, and established in, RŽunion (Aubert and Quilici 

1984), Mauritius (Quilici 1989) and Taiwan (Chiu et al. 1988). 

T. radiata was found to be the dominant parasitoid of D. citri on Murraya 

paniculata in Fujian, comprising 62.6% of all parasitoids and 

hyperparasitoids emerging. The second in abundance was the 

hyperparasitoid Tetrastichus sp., most of which were bred from T. radiata, 

an average of 21.8% hyperparasitisation, rising to a maximum of 87.9%, 

whereas the other primary parasitoid D. aligarhensis was hyperparasitised to 

an average of 34.1% (Tang 1989). 

The T. radiata female oviposits ventrally between the thorax and 

abdomen of the nymph, preferably of the 5th instar, and its fully grown larva 

spins silk to attach itself and its host to the plant substrate. The D. citri 

mummy parasitised by T. radiata has a dark brown, flattened body and the


parasitoid pupa remains external to, and on the ventral surface of, the host. 

The adult wasp emerges via a hole cut through the thorax of the host (Qing 

1990; Tang and Huang 1991). Under favourable conditions, parasitisation 

can exceed 90%, as in India (Husain and Nath 1927) and also in RŽunion, 

Nepal and Taiwan (Quilici and Fauvergue 1990). 

Male T. radiata are capable of multiple matings, but females usually 

mate only once. The egg to adult period was 11.4 days (egg 1.9, larva 4.0, 

prepupa 0.6, pupa 4.9 days), females lived 23.6 days and males lived 14.8 

days (Chien et al. 1991a,b; Chu and Chien 1991). Fauvergue and Quilici 

(1991) report reduction of the duration of immature stages with increasing 

temperature from 17 days at 20¡C to 8 days at 30¡C. Adult females lived 37 

days at 20¡C and 8 days at 35¡C. Females kill some 80% of D. citri hosts by 

parasitisation and 20% by host feeding. When 40 psyllids were presented per 

day a female killed 513 psyllids in a lifetime. At an optimum temperature of 

25¡C, 24, 5th instar nymphs were killed per day (Chien et al. 1993). Adult 

parasitoids can be cold stored at 8¡C for between 46 and 60 days (Chien et al. 

1993). Oosorption occurred when hosts were unavailable. This extended the 

reproductive period, but diminished the total number of eggs laid (Chien et 

al. 1994b). Feeding by females on the honeydew produced by the host and on 

host haemolymph provides nutrients for egg production. The parasitoid fed 

on the exudate of 28% of host eggs parasitised (Chien et al. 1994a). The 

optimal host density over the entire T. radiata lifetime was found to be 2 to 8 

per day, of which 90 to 94% were utilised. For the peak oviposition period, 

optimal density was 2 to 20, of which 87 to 90% were utilised (Chien et al. 

1995). 

The sex ratio of T. radiata is 1:3 in favour of females. Unmated females 

give rise only to male offspring. Oviposition occurs on 3rd, 4th and 5th instar 

nymphs and there is discrimination against ovipositing in nymphs 

containing older D. aligarhensis larvae. The average number of offspring is 

reported as 134 with 6.5 eggs laid per day (Tang and Huang 1991). 

Observations in China indicate that T. radiata is more affected by low 

temperatures than D. citri. Thus T. radiata breeds more effectively in 

Xiamen, where the lowest winter temperature is 3.9¡C, than in Fuzhou, 

where overwintering is jeopardised by lowest minimum temperatures of 

Ð2.5¡C (Aubert 1990).


Tetrastichus sp. Hym.: Eulophidae 

This undescribed species is an important hyperparasitoid of Tamarixia 

radiata in China. Average hyperparasitisation amounted to nearly 25%, with 

a maximum of 87.9%. The genus Tetrastichus contains more than 150 

species attacking a wide variety of hosts. 

ARACHNIDA 

Marpissa tigrina: Salticidae 

The number of D. citri consumed daily by an individual spider increased 

with an increase in available prey up to 40. Further increases in prey 

numbers reduced predation. The results suggest that M. tigrina is a highly 

efficient predator of D. citri (Sanda 1991). 

Comments 

Diaphorina citri and its two primary parasitoids, Diaphorencyrtus 

aligarhensis and Tamarixia radiata, are (especially the first two species) 

very widespread in Southeast Asia. In these countries the prospects for 

biological control are unpromising, although the careful timing of least 

harmful, but still effective, insecticides (or, preferably, special petroleum 

oils) would favour a build up of the parasitoids. The role played by hedges 

and other plantings of the common, favoured host, jasmin orange, Murraya 

paniculata in encouraging either D. citri or its parasitoids is worthy of 

careful investigation, for this may differ widely according to the insecticidal 

treatments in the nearby citrus plantings. Overhead irrigation to reduce 

numbers of eggs and young nymphs during periods of growth flushes is 

probably uneconomical in most situations, but is possibly a factor to 

consider in a pest management approach. 

Although D. aligarhensis appears to be a less effective parasitoid than 

T. radiata, it still may contribute useful suppression of D. citri where it can 

be introduced without encountering hyperparasitoids. 

In contrast to much of Southeast Asia, the prospects for successful 

biological control of D. citri appear to be promising for countries that have 

been recently invaded, particularly if there are few or no hyperparasitoids 

already present that are capable of attacking T. radiata and/or 

D. aligarhensis. In this context it may be valuable to explore the situation in 

Irian Jaya and Timor where D. citri has been recorded only recently. 

Successful biological control there may slow the spread of D. citri into 

Papua New Guinea, Australia and the oceanic Pacific. Brazil has a range of 

native psyllids that are attacked by Tamarixia spp. so it is possible that there 

are already hyperparasitoids present that would attack T. radiata were it to 

be introduced.


Since Tamarixia leucaenae attacks both Heteropsylla cubana and 

H. spinulosa, it would be valuble to know whether Tamarixia radiata will 

also attack Heteropsylla spinulosa. This is the psyllid that has been 

successfully introduced to Papua New Guinea, Australia and some oceanic 

Pacific countries for the biological control of creeping, sensitive plant, 

Mimosa invisa. If it does attack H. spinulosa, there would clearly be a 

conflict of interest between biological control of D. citri and of M. invisa, 

were Heteropsylla spinulosa to be considered for the latter. However, at 

least some species of Tamarixia are satisfactorily host restricted and it is 

quite possible that T. radiata is one of them.

4.8 Dysdercus cingulatus 

India 

Myanmar 

+ 

20° 

Laos 

+ 

0° 

20° 

China 

+ 

Thailand 

+ 

Cambodia 

P 

Vietnam 

++ 

P 

+ Brunei 

Malaysia 

+ 

Singapore 

++ 

Indonesia 

Taiwan 

++ 

Philippines 

Australia 

Papua 

New Guinea 

P 

135 

Dysdercus cingulatus is native to the Southeast Asian region. 

No parasitoids of the cotton stainer are known and surprisingly few predators are 

reported. No effective parasitoids of the many Dysdercus species that occur in other parts 

of the world are known. On present knowledge, therefore, the prospects for classical 

biological control of this bug would appear to be very remote. 

20° 

0° 

20°


Dysdercus cingulatus (Fabricius) 

Rating 

Origin 

Distribution 

Biology 

Hemiptera, Pyrrhocoridae 

cotton stainer, red cotton bug, red seed bug 


++ Viet, Indo, Phil 

11 + Myan, Thai, Laos, 

Msia, Sing 

+ 

P Brun P PNG, Sol Is, Van, Marianas, 

Carolines, N Cal. 

Southeast Asia. 

Waterhouse (1993a) was in error in accepting Kalshoven (1981), who listed 

the distribution of D. cingulatus as Ôwidespread from the Mediterranean to 

AustraliaÕ. In fact, it occurs from north eastern India through Bangladesh to 

all Southeast Asian countries, southern China, southern Japan, Irian Jaya, 

Papua New Guinea, eastern Australia, Saipan, Palau, Pohnpei, Yap, 

Solomon Is., Vanuatu and New Caledonia (CIE 1985). There are some 50 

species of Dysdercus, 

some of which appear to be native to the African, 

Ethiopian, Southern Asian and American regions respectively (Freeman 

1947). D. cingulatus is not recorded from Africa, Europe, Western Asia or 

the Americas. 

Eggs are usually laid singly in batches of about 100 (range 25 to 112) in 

small depressions in the soil under the host plant. They are camouflaged with 

soil particles or other debris. Some 80% of the eggs hatch in about 6 days if 

there is the essential high humidity. The optimum hatch occurs at 30¡C and 

80% RH. There are 5 nymphal instars which take 25 to 27 days to complete 

and the oviposition to adult period is thus 31 to 33 days. The first instar 

nymphs do not feed. Later instars suck sap with a preference for pods and 

seeds. All stages are gregarious. The male:female ratio is 3:2 and mating 

takes place readily and repeatedly, pairs often remaining in copula for 2 to 4 

days, during which they move and feed (Srivastava and Bahadur 1958; 

Thomas 1966; Ahmad and Aziz 1982, 1983, Farine and Lobreau 1984, 

Siddiqi 1985, 1987; Khoo et al. 1991).

Host plants 

Damage 

4.8 


137 

D. cingulatus is one of the commonest insects in cultivated land in 

Indonesia. Adults range from 11 to 17 mm in length and are orange and black 

in colour, with a characteristic white band on the pronotum and single large 

black spots on each orange forewing. 

Both adults and nymphs produce a complex mixture of compounds as a 

defensive secretion (Farine et al. 1992, 1993) and females produce a sex 

pheromone from glands in the thorax (Siddiqi 1988; Siddiqi and Khan 

1982). 

The principal host plants of D. cingulatus are in the families Malvaceae and 

Bombacaceae and include cotton, kapok, okra and rosella. It was the most 

important pest of cotton in the coastal districts of Malaysia in the early days 

and also fed on the seeds of kapok, rosella, hibiscus, hemp, okra and other 

malvaceous plants (Jack and Sands 1922; Dresner 1955). In a host 

preference test in Malaysia, the following order was recorded kapok > okra > 

urena > maize > sorghum (Chong 1975). It has also been recorded on wheat 

(Srivastava and Gupta 1971), pearl millet, Pennisetum glaucum 

(= P. americanum) 

(Ahmad 1979) and a range of weeds. 

Like other species of Dysdercus throughout the world D. cingulatus is most 

important as a pest of cotton. The sap removed and the fungus introduced 

into the punctures caused when feeding on the developing cotton bolls 

causes staining of the lint, giving rise to one of its common names. 


Very little has been published on the natural enemies of D. cingulatus and no 

parasitoids are known. In view of the large number of parasitoids of Nezara 

viridula eggs (Table 4.12.1), it seems strange that none are recorded from 

eggs of the cotton stainer. There appear to be no statements in the literature 

that they have been looked for unsuccessfully. 

The few records retrieved of natural enemies of D. cingulatus are shown 

in Table 4.8.1. The pyrrhocorid predator Antilochus coquebertii has been 

reported attacking the cotton stainer in India and Malaysia (Yunus & Ho 

1980; Zaidi 1985). It is believed to inject saliva containing proteolytic 

enzymes into eggs, nymphs or adults before sucking out the liquified 

contents. In the Philippines an ectoparasitic mite, Hemipterotarseius sp. was 

found on the pronotum of adult bugs, two species of spiders were observed 

attacking bugs and a nematode was found in the abdomen of a female. No


Comment 

Table 4.8.1 

egg parasitoids were found (Encarnacion 1970). Singh and Bardhan (1974) 

showed that D. cingulatus was moderately susceptible to the DD 136 strain 

of the nematode Steinernema carcocapsae. 

A superficial review of the literature on the natural enemies of other 

species of Dysdercus revealed two tachinid parasitoids, one of each in South 

America and Africa and three asassin bug predators in Africa but no egg 

parasitoids (Table 4.8.2). 

Too little is known about the natural enemies of D. cingulatus to indicate 

whether there are any prospects for classical biological control or for 

manipulating them where they already occur. Since D. cingulatus now 

appears to have evolved in the Southeast Asian region and since effective 

natural enemies of related species elsewhere are not known, the prospects 

for its classical biological control would appear to be remote. 

Natural enemies of Dysdercus cingulatus 

Enemy 

INSECTA 

HEMIPTERA 


PYRRHOCORIDAE 

Antilochus coquebertii 

India 

Malaysia 

Zaidi 1985 

Thomas 1966; Yunus & Ho 1980 

ACARINA 

Hemipterotarseius sp. 

ARACHNIDA 

Philippines Encarnacion 1970 

Spider 1 Philippines Encarnacion 1970 

Spider 2 

NEMATODA 


Species 1 

FUNGI 


Aspergillus flavus 

India Kshemkalyani et al. 1989

Table 4.8.2 

4.8 

Natural enemies of Dysdercus spp. 


Enemy 

HEMIPTERA 

REDUVIIDAE 


Phonoctonus nigrofasciatus Zimbabwe Sweeney 1960 

Phonoctonus subimpictus Ivory Coast Galichet 1956 

Phonoctonus sp. Mozambique Barbosa 1950 

DIPTERA 

TACHINIDAE 

Acaulona brasiliana 

Bogosia helva 

Argentina Blanchard 1966 

Ivory Coast Galichet 1956 

139

4.9 Dysmicoccus brevipes 

India 

20° 

0° 

20° 

Myanmar 

Laos 

China 

+ 

Thailand 

P 

Cambodia 

+ 

Vietnam 

+++ 

P 

++ Brunei 

Malaysia 

Singapore 

++ 

Indonesia 

Taiwan 

+ 

++ 

Philippines 

Australia 

Papua 

New Guinea 

+ 

141 

The pineapple mealybug, Dysmicoccus brevipes, 

is of Central or South American 

origin. It occurs in both parthenogenetic and bisexual forms, the females of which are 

morphologically indistinguishable, although they may possibly prove to be distinct species. 

The closely-related, bisexual, D. neobrevipes also occurs on pineapple. 

Attempts have been made by several countries to establish natural enemies for 

biological control, but none has had any great success without the control of attendant 

ants. When ants were controlled the mealybug was no longer a problem. Ant control is less 

likely to be an answer to the problem in the absence of suitable natural enemies. It is 

desirable, therefore, to establish appropriate natural enemies in anticipation of effective 

ant control, an aspect which is now being actively investigated in Hawaii. 

20° 

0° 

20°


Dysmicoccus brevipes (Cockerell) 

Rating 

Origin 

Distribution 

Taxonomy 

Hemiptera, Pseudococcidae 

pineapple mealybug 


+++ Viet +++ Cook Is, Guam 

10 ++ Msia, Indo, Phil 17 ++ FSM, Niue, Van 

+ Camb + + Kiri, N Cal, PNG, Sam, 

Sol Is 

P Brun, Thai P Fiji, Fr P, A Sam. 

Tok, Tong, Tuv 

Carter (1935) considered D. brevipes to be native to South America, 

although Ferris (1950) believed it to be of North American origin. The 

pineapple plant ( Ananas comosus) 

is thought to be native to South America 

and has been known in Central America since pre-Columbian times. 

Although the pineapple mealybug is widely polyphagous, if it evolved in 

association with pineapple plants, it would appear to be logical to assign its 

origin to Central and/or South America. 

D. brevipes is one of the most widespread mealybugs, occurring throughout 

the tropics and in many temperate areas, especially those where pineapples 

are grown (Williams and Watson 1988). These include tropical Africa, 

Mauritius, tropical Asia, Southeast Asia, Taiwan, Australia, Pacific islands 

(including Hawaii), southern USA (Florida, Louisiana) West Indies and 

Central and South America (Bartlett in Clausen 1978). 

Dysmicoccus brevipes was earlier known as Pseudococcus brevipes, 

but its 

genus was changed by Ferris (1950). Before 1959 it was confused with a 

similar mealybug (often on the same host plants), which was described by 

Beardsley (1959) as Dysmicoccus neobrevipes. 

Both species occur in 

Hawaii, where D. brevipes is parthenogenetic and the females are pink, 

whereas D. neobrevipes is bisexual and females are grey in colour. 

Parthenogenetic D. brevipes is known also from Jamaica and West Africa 

(Beardsley 1965).

Biology 

4.9 


143 

In some countries (Ivory Coast, Madagascar, Dominican Republic, 

Martinique, Malaysia) both sexes of D. brevipes occur (Beardsley 1965; 

Lim 1973). D. neobrevipes is known from many countries in Central and 

South America and probably also originated there (Williams and Watson 

1988). It is known from Mexico, Jamaica, American Samoa and Samoa, 

Cook Is, Kiribati and Guam. In Southeast Asia it occurs in Malaysia, the 

Philippines and Thailand (where there have been recent serious outbreaks 

(Beardsley 1965; Rohrback et al. 1988; Williams and Watson 1988). 

D. brevipes females are broadly oval to rotund in shape, pinkish in colour, 

and have a thick waxy covering with short conical waxy projections 

(Kalshoven 1981). In Hawaii, D. brevipes is parthenogenetic and 

ovoviviparous (i.e. it produces its young alive). About 250 crawlers are 

produced per female over a 3 to 4 week period and take some 34 days to 

mature. Females start producing young about 25 days after the third moult 

(Ito 1938). 

In peninsular Malaysia the bisexual form is widespread but the 

parthenogenetic form was not found (Lim 1973). In the male, there are two 

nymphal, one prepupal and one pupal instar of 10, 6, 3 and 4 days duration 

respectively. Adult males live for 1 to 3 days, whereas adult females live 17 

to 49 days. The female has 3 nymphal instars, lasting 10, 7 and 7 days 

respectively. A female produces 19 to 137 offspring with a sex ratio of 1:1 

(Lim 1973). The bisexual form in Malaysia has a 10 day shorter life cycle 

than the parthenogenetic form in Hawaii and might have as many as 9 

generations a year (Lim 1973). The bisexual form of D. brevipes is capable 

of producing green spotting on pineapple leaves, whereas the 

parthenogenetic form is not (Beardsley 1965). 

In India, Ghose (1983) studied the parthenogenetic form of D. brevipes 

at 30¡C and 60Ð66% R.H. The nymphs completed their development in 19 

days and, after a pre-oviposition period of 16 days, produced an average of 

240 young over a period of 40 days. 

D. brevipes occurs mainly on the underground parts of the pineapple 

stem (where the stem is covered by leaf bases) and on the roots. Leaves or 

fruit are less heavily infested, except on weak plants. However the bisexual 

form also infests the crown of the pineapple plant (Rohrbach et al. 1988). 

By comparison, D. neobrevipes, 

which is always bisexual, is found only 

on the aerial parts of the pineapple plant Ñ leaves, stems, aerial roots, 

flowers and fruit clusters. Unlike D. brevipes, 

it does not infest grasses . In 

Hawaii, only D. neobrevipes causes green spotting of pineapple leaves 

(Beardsley 1959).


Hosts 

Damage 

The pineapple mealybug is generally attended by ants seeking 

honeydew. They not only protect it against natural enemies but also assist in 

dispersal by transporting it to new plants. Where appropriate natural enemies 

are present, but attendant ants are not, the mealybug is no longer a problem 

(Beardsley et al. 1982). The identity of the ants varies from place to place, 

although 3 very widespread species are often involved, the bigheaded ant, 

Pheidole megacephala, 

the Argentine ant, Iridomyrmex humilis, 

and the fire 

ant Solenopsis geminata. 

The gradual invasion of new pineapple plantings 

by ants is accompanied by progressive outbreaks of mealybugs, so ant 

control is essential. It is interesting that mated queens of P. megacephala 

must, following the nuptial flight, rejoin established colonies to survive. 

Thus, invasion into new territory is accomplished by extension of existing 

nests, a feature that is of importance in controlling the big-headed ant 

(Beardsley et al. 1982). 

D. brevipes occurs widely on its preferred host, pineapple, wherever this is 

grown in moist tropical or subtropical areas, but it can be found on almost 

any kind of plant and is sometimes a pest of sugarcane and bananas. It also 

occurs on areca palm, coffee, groundnut, oil palm, rice, sisal, soybean, 

Pandanus palm and a range of grasses and weeds (Clausen 1978; Kalshoven 

1981; Khoo et al. 1991). 

In Hawaii, through their feeding, both D. brevipes and D. neobrevipes 

produce symptoms of toxicosis on pineapple, including stunting, reddening 

and wilting of young plants, due to what is now held to be a virus, and termed 

pineapple mealybug wilt. D. neobrevipes, 

but not the parthenogenetic form 

of D. brevipes, also produces a green spotting on the leaves. Where the 

bisexual form of D. brevipes occurs elsewhere it is capable of producing 

green spotting. 

Unless collected by ants, honeydew produced by the mealybugs, leads to 

the massive growth of sooty moulds which reduces photosynthesis, affects 

sales of fruit and attracts Carpophilus spp. beetles, which contaminate 

canned fruit. Large colonies of D. brevipes in leaf sheaths near the roots of 

sugarcane result in poor growth and, when on groundnuts, cause the seed 

pods to become discoloured.


4.9 


145 

A number of coccinellid predators are recorded attacking D. brevipes (Table 

4.9.1) and it is possible that some of these have a sufficiently narrow host 

range to be considered for introduction. However, it is more likely that the 

dipterous and lepidopterous predators will be more specific. The encyrtid 

parasitoids would appear to be even more promising and it seems that the full 

range of species attacking D. brevipes in Central and South America has not 

yet been identified. 


HAWAII 

A number of natural enemies, mainly encyrtid parasitoids and coccinellid 

and dipterous predators have been successfully introduced, in particular into 

Hawaii, Puerto Rico and the Philippines (Table 4.9.2). Details are provided 

in the country accounts that follow. It will become clear that a substantial 

degree of control of D. brevipes can be achieved in the absence of ants, 

which clearly protect the mealybugs against parasitoids and predators. 

The pineapple mealybug has, for many years, constituted the most serious 

insect problem of the pineapple industry (Carter 1932; Beardsley 1959) and 

it is also a minor pest of sugarcane and bananas. An encyrtid wasp 

Euryrophalus schwarzi (= E. pretiosa) 

was reared from D. brevipes 

collected from sugarcane (Beardsley 1959). 

In the early 1920s several natural enemies were introduced from Mexico 

and Panama, but none became established (Rohrbach et al. 1988). Anagyrus 

ananatis and Hambletonia pseudococcina from Central America and Brazil 

were established in 1935Ð36 and were effective in Maui where the dominant 

ant was the crazy ant Paratrechina longicornis. 

Parasitisation was high and 

pineapple wilt quite severe (Carter 1945). Other introductions known to 

have become established are an encyrtid parasitoid ( Euryrhopalus 

propinquus), 

a cecidomyiid predator ( Vincentodiplosus pseudococci), 

and 

two less effective coccinellid predators ( Scymnus (= Nephus) 

bilucenarius 

and Scymnus uncinatus) 

(Lai and Funasaki 1986). 

Overall, although the biological control of the pineapple mealybug has 

not been completely successful, a considerable reduction in abundance has 

resulted from the combined action of the cecidomyiid predator 

Vincentodiplosis and the encyrtid parasitoids Anagyrus ananatis and 

Hambletonia pseudococcina. 

They are highly effective only where ants are 

adequately controlled (Clausen 1978). Coccinellids are important for short 

periods, particularly in the middle of large plantings, where the absence of 

ants renders D. brevipes exposed to attack (Carter 1935, 1944).

Table 4.9.1 

Natural enemies of Dysmicoccus brevipes 

Country References 

HEMIPTERA 

DIASPIDIDAE 

Diaspis bromeliae 

ORTHOPTERA 

Mauritius Jepson 1939a 

Conocephalus saltator 

NEUROPTERA 

CHRYSOPIDAE 

Hawaii Carter 1935 

Chrysopa irregularis 


Chrysopa ramburi 


Chrysopa sp. 

COLEOPTERA 

COCCINELLIDAE 


Brachycantha sp. Guatemala Carter 1935 

Cryptolaemus montrouzieri 

Hawaii 


Fiji, New Caledonia 

Williams & Watson 1988 

Cryptolaemus sp. Fiji, New Caledonia Williams & Watson 1988 

Rhizobius ventralis 

Hawaii, New Caledonia Williams & Watson 1988 

Scymnus apiciflavus 

Malaysia Yunus & Ho 1980 

Scymnus bilucenarius 

Guatemala Carter 1935 

Scymnus mauritiusi 

Mauritius Jepson 1939a 

Scymnus sp. Fiji, New Caledonia Cohic 1958 

Sticholotis quatrosignata 


Coccinellids 

Other species in introductions listed in table 4.9.2 

Taiwan Takahashi 1939 


Table 4.9.1 (contÕd) Natural enemies of Dysmicoccus brevipes 

DIPTERA 


CECIDOMYIIDAE 

Schizobremia formosana Taiwan Takahashi 1939 

Cecidomyiid sp. 1 Guatemala Carter 1935 

Cecidomyiid sp. 2 Mauritius Jepson 1939a 

Cecidomyiid sp. 3 Puerto Rico Plank & Smith 1940 

DROSOPHILIDAE 

Gitonides perspicax Mauritius Jepson 1939a 

Pseudiastata nebulosa Guatemala Carter 1935 

LEPIDOPTERA 

PYRALIDAE 

Species 1 

TINEIDAE 

Puerto Rico Plank & Smith 1940 

Drosica abjectella South Africa BŸttiker 1957 

Species 1 

HYMENOPTERA 

ENCYRTIDAE 

Puerto Rico Plank & Smith 1940 

Aenasius acuminatus Trinidad Kerrich 1967 

Aenasius theobromae Trinidad Kerrich 1953 

Anagyrus ananatis Brazil Carter 1937; 

Gabriel et al. 1982 

Anagyrus sp. Brazil Compere 1936 

Encyrtid sp. 1 Brazil Compere 1936 

Euryrhopalus propinquus Brazil, Guyana, Hawaii Kerrich 1967 

4.9 


147

Table 4.9.1 (contÕd) Natural enemies of Dysmicoccus brevipes 

HYMENOPTERA 

ENCYRTIDAE (contÕd) 

Euryrhopalus schwarzi (= E. pretiosa) Guatemala 

Hawaii 


Clausen 1978 

Beardsley 1959 

Hambletonia pseudococcina Brazil, Colombia, Venezuela Carter 1937 

Leptomastix dactylopii California Clausen 1978 

Pseudaphycus angustifrons Cuba Gahan 1946 

Pseudaphycus dysmicocci Trinidad Clausen 1978 

Pseusaphycus sp. Brazil Clausen 1978 

Thysanus niger 

CHALCIDIDAE 

Puerto Rico Bartlett 1945 

Species 1 Guatemala Carter 1935 

Species 2 

UNIDENTIFIED FAMILY 

Guatemala Carter 1935 

7 species 

ARACHNIDA 


spiders Hawaii Carter 1944 


Table 4.9.2 Introductions for the biological control of Dysmicoccus brevipes 

Country From Year Result Reference 

HYMENOPTERA 

ENCYRTIDAE 

Aenasius colombiensis Colombia 1935 Ð Lai & Funasaki 1986 

Aenasius sp. Panama 1931 Ð Lai & Funasaki 1986 

Anagyrus ananatis 

Hawaii Brazil 1934Ð35 + Carter 1937 

(= Anagyrus coccidivorus) 

Peurto Rico Brazil 1937Ð38 Ð Bartlett 1939, 1943 

via Hawaii 

Clausen 1978 

Anagyrus kivuensis California 1946 Ð Lai & Funasaki 1986 

Euryrhopalus propinquus Hawaii British 

Guiana 

1935 + Lai & Funasaki 1986 

Euryrhopalus schwarzi Hawaii Guatemala 1935 + Clausen 1978 

Hambletonia pseudococcina Hawaii Brazil 1935Ð36 Ð Carter 1937; Clausen 1978 

Colombia 1935Ð36 + Carter 1937; Clausen 1978 

Venezuela 1935Ð36 + Carter 1937 

Jamaica Hawaii 

1936 Ð Clausen 1978 

Puerto Rico Brazil via 

Hawaii 

1937Ð38 + Bartlett 1939 

Florida Puerto Rico 1944 + Annand 1945; Clausen 1956 

Leptomastix dactylopii Hawaii California ? Clausen 1978 

Pseudaphycus dysmicocci Hawaii Trinidad 1958 ? Clausen 1978 

Pseudaphycus sp. Hawaii Brazil 1946 ? Clausen 1978 

Zaplatycerus fullawayi 

PLATYGASTERIDAE 

Mexico 1930 Ð Lai & Funasaki 1986 

Allotropa sp. Panama 1931 Ð Lai & Funasaki 1986 

4.9 Dysmicoccus brevipes 149

Table 4.9.2 (contÕd) Introductions for the biological control of Dysmicoccus brevipes 

COLEOPTERA 

COCCINELLIDAE 

Cleothera sp. Panama 1931 Ð Lai & Funasaki 1986 

Cryptolaemus montrouzieri Mauritius 

Easter Is 

(Chile) 

S. Africa 1938Ð39 Ð 

+ 

Jepson 1939b, 

Moutia & Mamet 1946 

Ripa et al. 1995 

Cryptolaemus sp. Taiwan ? Sakimura 1935 

Diomus sp. Jamaica 

Panama 


Hawaii 1939 

1931 

Ð 

Ð 

Lai & Funasaki 1986 


Hyperaspis albicollis Panama 1924 Ð Lai & Funasaki 1986 

Hyperaspis c-nigrum Brazil 1935 Ð Lai & Funasaki 1986 

Hyperaspis silvestri Philippines Hawaii 1931 + Clausen 1978 

Hyperaspis 12 ´ spp. Hawaii various ? Clausen 1978 

Hyperaspis sp. Jamaica Hawaii 1939 Ð Clausen 1978 

Scymnus (=Diomus) margipallens Philippines Hawaii 1931 + Clausen 1978 

Scymnus (=Nephus) bilucernarius Hawaii 

Mexico 1930 + 



Scymnus pictus Panama 1924 Ð Lai & Funasaki 1986 

Scymnus quadrivittatus California 1948 Ð Lai & Funasaki 1986 

Scymnus uncinatus Hawaii Mexico, 

Panama 

1922 + Lai & Funasaki 1986 

Scymnus 6 ´ spp. Hawaii various ? Clausen 1978 

Scymnus sp. Taiwan Saipan ? Sakimura 1935 


Table 4.9.2 (contÕd) Introductions for the biological control of Dysmicoccus brevipes 

DIPTERA 


CECIDOMYIIDAE 

Cecidomyiid sp. Panama 1931 Ð Lai & Funasaki 1986 

Cleodiplosis koebelei Philippines Hawaii 1931 + Clausen 1978 

Dicrodiplosis guatemalensis Hawaii Guatemala 1935 + Clausen 1978 

Vincentodiplosis (= Lobodiplosis) pseudococci Hawaii Mexico 1930 + Clausen 1978 

DROSOPHILIDAE 

Pseudiastata nebulosa Hawaii Guatemala 1924 

1932 

Ð 

Ð 

Carter 1935 

Carter 1935 

Pseudiastata pseudococcivora Panama 1931, 1951 Ð Lai & Funasaki 1986 

4.9 Dysmicoccus brevipes 151


BRAZIL 

Bisexual D. brevipes is present wherever pineapples are grown, but it is not 

recorded whether the parthenogenetic strain also occurs. Mealybug wilt was 

rare in 1946 and large mealybug colonies uncommon and always covered 

with soil mounds built by Solenopsis sp. ants. Sparse green spotting was 

general on the leaves of these plants. Natural enemies were numerous and 

included Anagyrus sp., Hambletonia pseudococcina and Pseudaphycus sp. 

The latter parasitised mealybugs on the aerial parts of the pineapple plant, so 

that large colonies were rare. It was never found on colonies under the soil 

surface. Its life cycle lasts 14 to 20 days and up to 6 individuals may emerge 

from a single host. Predators, mainly coccinellids, were generally present 

and attacked the underground mealybug colonies (Carter 1949). 

COOK IS 

Carter (1973) reported that mealybug wilt of pineapple was a serious threat 

to the newly-developing pineapple industry on Atui and Mangaia. 

D. brevipes was present, but not D. neobrevipes, and the mealybug was 

attended by Pheidole megacephala. 

FIJI 

D. brevipes is the main pest on pineapple and causes pineapple wiltÑthe 

only record in the Pacific outside Hawaii of this condition. It is also a minor 

pest of sugarcane. It is controlled to some degree by the coccinellids 

Cryptolaemus sp., C. montrouzieri and Scymnus sp. (Lever 1945). Three 

chrysopids were predators, including Chrysopa ramburi and C. irregularis 

(Lever 1940). 

GUATEMALA 

Two coccinellid predators of D. brevipes were reported by Carter (1935), 

Scymnus bilucenarius was widespread except in highest elevations, but it 

apparently exerted little control; and Brachycantha sp. which was 

uncommon. A drosophilid fly, Pseudiastata nebulosa, which was heavily 

parasitised by two chalcidid wasps, was found and frequently in large 

numbers. It was regarded as a promising species for biological control by 

Carter (1935), who introduced it to Hawaii in 1932, but the colony died out. 

Cecidomyiid predators were very common in the lowlands and occurred 

occasionally in the highlands. They attacked large mealybug colonies on 

fruit, but were apparently a minor control factor. No hymenopterous 

parasitoids were discovered. 

GOLD COAST 

The parasitoid Pseudaphycus angelicus was reared on D. brevipes in the 

laboratory (Anon. 1953) as also was Anagyrus ananatis from California 

(Anon. 1957). Both were released in 1953Ð54 and the former became 

established. Scymnus sordidus was also introduced from California, reared 

on D. brevipes and released, but establishment is not reported (Anon. 1957).

4.9 Dysmicoccus brevipes 153 

INDONESIA 

Kalshoven (1981) reported that D. brevipes is attended by the ant, 

Monomorium sp. 

IVORY COAST 

D. brevipes is bisexual with a sex ratio usually of 2 males to 1 female. Ants 

attending the pineapple mealybug were species of Camponotus, 

Crematogaster and Pheidole (Re‡l 1959). 

JAMAICA 

Here and throughout Central America D. brevipes colonies of any size were 

invariably attended by Solenopsis ants. Where Solenopsis was not present 

mealybug colonies were rare and small (Carter 1935). 

MALAYSIA 

The pineapple mealybug D. brevipes is the most serious insect pest of 

pineapple in peninsular Malaysia. Infected plants become stunted and 

reddish and eventually wilt. Fruit are small and unsuitable for canning. In 

addition to wilting, the mealybug causes green spotting of the leaves, which 

is not of economic importance (Khoo et al. 1991). 

The bisexual form of D. brevipes has been studied in some detail by Lim 

(1972). It was the only form found in 14 pineapple areas visited in Johore 

and Selangor. The bisexual form had a life cycle 10 day shorter than the 

parthenogenetic form in Hawaii, although it was less prolific. 

MAURITIUS 

D. brevipes was first reported as a major pest of pineapples in 1933, probably 

having been introduced on pineapple suckers from Hawaii about 1931. It 

was attacked by three native predators, the coccinellid Scymnus mauritiusi, 

the drosophilid fly Gitonides perspicax, and the pineapple scale, Diaspis 

bromeliae (Jepson 1939b) but they produced little impact. The coccinellid 

Cryptolaemus montrouzieri was introduced from South Africa and liberated, 

but did not become established (Moutia and Mamet 1946). The mealybug is 

attended by Pheidole megacepahala, Solenopsis geminata and 

Technomyrmex detorquens (Jepson and Wieke 1939). 


D. brevipes is recorded attacking taro where it is subject to predation by the 

coccinellids Cryptolaemus affinis, C. montrouzieri and C. wallacii (Shaw et 

al. 1979). 

PHILIPPINES 

Two strains (grey and pink) of the pineapple mealybug are present and it is 

suggested that they may have been introduced with planting material from 

Hawaii, in which case they would represent D. neobrevipes and D. brevipes 

respectively. The grey strain produces green spotting of the leaves, whereas 

the pink strain produces only chlorotic spots. Some pineapple cultivars can 

be seriously damaged by pineapple wilt which is caused by both strains. Two


species of ants are almost invariably in attendance on the mealybugs, namely 

Solenopsis geminata and Pheidole megacephala (Serrano 1934). Three 

predators from Hawaii were established in 1931 (Cleodiplosis koebelei: 

Cecidomyiidae, Scymnus margipallens and Hyperaspis silvestri: both 

Coccinellidae), but there is no information on their effectiveness (Clausen 

1978). 

PUERTO RICO 

D. brevipes is the most serious pest of pineapples, attacking the roots, leaves 

and fruits of all varieties grown. Anagyrus ananatis was imported in 1936 

from Brazil and both A. ananatis and Hambletonia pseudococcina in 1937 

from Hawaii where they had been introduced from Brazil and Venezuela 

respectively. Both were released in 1937 and 1938. Only half grown or older 

hosts were attacked by H. pseudococcina, the life cycle from oviposition to 

adult emergence taking 24 to 30 days. In one instance, 3 parasitoids emerged 

from the same host. Frequent recoveries were made from the field. 

Development from egg to adult took 19 to 21 days for A. ananatis and the sex 

ratio was 1:1 (Bartlett 1939). Recoveries of this species were reported later 

(Bartlett 1943). The larvae of a tineid moth, a pyralid moth and of a 

cecidomyiid fly were found living in the waxy secretions around large 

groups of mealybugs and were thought to be predators. Three species of ants, 

including Solenopsis geminata were frequently observed carrying young 

mealybugs around. There do not appear to be any recent reports of the 

effectiveness of the introduced parasitoids (Plank and Smith 1940). 

SOUTH AFRICA 

Larvae of the moth Drosica abjectella were observed preying on D. brevipes 

on pineapple in the Transvaal. The number of moth larvae and pupae per 

pineapple plant varied from 1 to 15 and these occurred in the leaf axils. Large 

nymphs and fully-fed mealybug females were preferred and moth larvae 

each consumed an average of 6.5 hosts in the laboratory. In winter, 

development of fourth instar D. abjectella took 15 to 26 days, the prepupal 

period 1 to 3 days and the pupal stage 35 to 45 days. The adults lived for 3 to 

6 days (BŸttiker 1957). 

SRI LANKA 

Both a bisexual and a parthenogenetic strain of D. brevipes are present. The 

former, which causes green spotting of pineapple leaves, occurs in the west 

and the latter in the Bibile area (Carter 1956). 

TAIWAN 

D. brevipes is widely distributed on pineapple up to about 750 m and also 

occurs on banana (Chiu and Cheng 1957). It appears that D. neobrevipes is 

also present. Natural enemies include the cecidomyiid predator 

Schizobremia formosana and also coccinellids, but these are less effective

TRINIDAD 

USA 

4.9 Dysmicoccus brevipes 155 

(Takahashi 1939). Cryptolaemus, imported for control of D. brevipes and 

other mealybugs, was only partially effective. A Scymnus sp., said to be 

effective against D. brevipes in Saipan, was introduced, but no further 

information is available (Sakimura 1935). The most abundant attendant ants 

were Pheidologeton diversus, Anoplolepis longipes and Camponotus 

friedae (Lee 1974). 

The encyrtid Pseudaphycus dysmicocci was reared as a solitary parasitoid of 

second instar female nymphs of D. brevipes on pineapple (Bennett 1955). 

D. brevipes was a common pest in southern Florida and Hambletonia 

pseudococcina was introduced from Puerto Rico and liberated in 1954. 

Although it became established, information on its abundance and 

effectiveness is not available. It was postulated that the widespread use of 

organic pesticides had probably reduced the parasitoid to very low levels 

(Clausen 1956). 


Anagyrus ananatis Hym.: Encyrtidae 

This moderately polyphagous wasp is widespread in Brazil, where it is 

known as a parasitoid of D. brevipes, but also parasitises, inter alia, rhodes 

grass scale (Antonina graminis) and citrus mealybug (Planococcus citri) 

(Gabriel et al. 1982). It was established in Hawaii (Carter 1937), where it 

completes a generation in about 20 days. 

Hambletonia pseudococcina Hym.: Encyrtidae 

This parasitoid occurs as a bisexual form on D. brevipes in Brazil and as a 

parthenogenetic one in Colombia and Venezuela. The bisexual form failed 

to reproduce on D. brevipes in Hawaii, but the parthenogenetic form did so 

successfully (Carter 1937). About 24 to 30 days is required for the life cycle 

under tropical outdoor conditions and up to 4 individuals may emerge from a 

single host (Compere 1936). In laboratory trials, H. pseudococcina showed 

a high degree of specificity for D. brevipes and did not oviposit in 8 closely 

related mealybug species. Of 3 additional mealybugs tested, it attempted 

oviposition only in an unidentified species from a grass (Clancy and Pollard 

1947). 

Pseudiastata nebulosa Dipt.: Drosophilidae 

This predator is native to Guatemala. It was introduced to Hawaii in 1924 but 

did not become established. Except in the highlands of Guatemala, it is 

frequently found in large numbers on a single plant, both above and below 

the soil line. It was again introduced in 1932, but the colony died out. This


was possibly due to its requirement for large numbers of hosts, since 7 larvae 

consumed over 100 medium to large sized hosts during the last half of their 

larval lives. In Guatemala it is highly parasitised by two species of chalcidid 

wasps. Carter (1935) regarded it as a promising species for biological 

control. 

Vincentodiplosis pseudococci Dipt.: Cecidomyiidae 

This midge is native to Mexico and was established in Hawaii in 1950. In 

neglected, weedy pineapple plantations in Hawaii, where the ant Pheidole 

sp. was less reliant on mealybug honeydew, the midge was sufficiently 

effective in controlling D. brevipes as to almost completely eliminate the 

mealybug from the fruit. Many fruit were covered with the old webs 

produced by midge larvae, but there were no live mealybugs. The midge is 

rarely found on leaves, but its larvae are commonly found attacking large 

mealybugs at the base of the fruit (Carter 1935, 1944). 

Comments 

It is very likely that D. brevipes evolved in South and/or Central America 

and there is, therefore, a prima facie case to consider it as a candidate for 

classical biological control in Southeast Asia and the Pacific. Indeed, there 

are 2 parasitoid species (Anagyrus ananatis and Hambletonia 

pseudococcina: both Eulophidae) and 2 predator species (Vicentodiplosis 

pseudococci: Cecidomyiidae and Pseudiasta nebulosa: Diastadidae) that are 

capable of reducing the mealybug to subeconomic levels. However, when 

any one or more of a number of ant species attends the mealybug it is largely 

protected from natural enemies and is able to build up to damaging numbers. 

In the absence of both ants and natural enemies the unharvested honeydew it 

produces leads to heavy growth of sooty moulds and there is transmission of 

pineapple mealybug wilt. Since apparently suitable natural enemies are 

available for introduction, the key to D. brevipes control is to deal with the 

attendant ants. There are several ant baits that have been used successfully 

for this purpose, but these are no longer registered for use in USA and, 

hence, cannot be recommended. No doubt suitable replacements will soon 

emerge. When extensive plantings of pineapples are made on areas where 

the soil has been worked to kill weeds, very few ant colonies survive. 

Recolonisation of the planted area occurs as colonies move along the rows 

towards the centre of the crop. One cultural method recommended to delay 

this spread is to plant several peripheral rows parallel to each boundary. Ants 

will then move along these, rather than into the crop and control measures 

can be concentrated on these rows (Rohrback et al. 1988). Promising results 

obtained with the integrated management of D. brevipes in Hawaii suggest 

that it would be well worth exploring similar methods elsewhere.

4.10 Hypothenemus hampei 

India 

20° 

0° 

20° 

Myanmar 

Laos 

+ 

China 

Thailand 

+ 

Cambodia 

P 

Vietnam 

++ 

+ 

++ Brunei 

Malaysia 

Singapore 

++ 

Indonesia 

Taiwan 

+++ 

Philippines 

Australia 

P 

Papua 

New Guinea 

157 

Hypothenemus hampei is native to Central Africa but has spread to most coffee 

producing countries in Central and South America, to Southeast Asia and to several 

Pacific countries. Significant coffee-growing areas not yet infested are Hawaii, Papua New 

Guinea, Vanuatu and Solomon Islands. 

It is a pest exclusively of coffee berries and does not damage the vegetative parts. It is 

difficult to control with chemicals and, although plantation management methods can 

reduce damage, the coffee berry borer remains an important pest. 

The most important natural enemies appear to be 3 parasitic wasps native to Africa, 

Cephalonomia stephanoderis, 

Phymastichus coffea and Prorops nasuta. 

The last of these 

has been established in Brazil and Colombia without its own natural enemies, but has not 

so far produced spectacular results. C. stephanoderis has been established recently in 

Colombia, Ecuador, Mexico and New Caledonia, but it is too early to evaluate its impact. 

Phymastichus coffea has not yet been established anywhere, but this is foreshadowed in 

Colombia. The fungus Beauveria bassiana shows early promise. A thorough study is in 

progress of the interactions of the parasites and other natural enemies of H. hampei and 

the influence on them of various components of the environment. Optimism has been 

expressed about the outcome of this program. 

20° 

0° 

20°


Hypothenemus hampei (Ferrari) 

Rating 

Origin 

Distribution 

Coleoptera: Scolytidae 

coffee berry borer 

Southeast Asia Southern and Western Pacific 

+++ Phil +++ N Cal 

12 ++ Viet, Msia, Indo 7 ++ Fiji, Fr P 

+ Thai, Laos, Brun 

P Camb P Pohnpei, Saipan 

This account updates the chapter on H. hampei in Waterhouse and Norris 

(1989) and the valuable review of Murphy and Moore (1990) in relation to 

prospects for biological control. 

FerrariÕs specimens, described in 1867 under the generic name 

Stephanoderes, 

were obtained from trade coffee beans in France. There 

appears to be no record of the country of origin of the material, but in 1867 

infested beans could only have come from Africa or Saudi Arabia, because 

Hypothenemus hampei did not obtain a footing on other continents until 

later. The seed used to establish Coffea arabica in Saudi Arabia was 

probably obtained from the Ethiopian highlands centuries ago. There the 

coffee berry borer is native, though scarce (Davidson 1967), but if it did not 

accompany the original seed it could easily have reached Saudi Arabia 

through Arabian-African commerce over the centuries. 

The wider range of parasitoids (3) in West Africa than in East Africa (2, 

with one shared with the West) suggests that H. hampei has been in the West 

for a very long time and may indeed have evolved there (L.O. Brun pers. 

comm.). 

This was given by CIE (1981) as: Africa (Angola, Benin, Burundi, 

Cameroon, Canary Is, Central African Republic, Chad, Congo, Ethiopia, 

Fernando Poo, Gabon, Ghana, Guinea, Ivory Coast, Kenya, Liberia, 

Malawi, Mozambique, Nigeria, Principe, Rio Muni, Rwanda, S‹o TomŽ, 

Senegal, Sierra Leone, Sudan, Tanzania, Togo, Uganda and Zimbabwe); 

Middle East (Saudi Arabia), Asia (Indonesia, Cambodia, Laos, Malaysia, 

Philippines, Sri Lanka, Thailand, Vietnam); Central America (Guatemala,

Biology 

4.10 


159 

Honduras, Greater West Indies); South America (Brazil, Peru, Surinam); 

Pacific (Caroline Is, Irian Jaya, Marianas Is, New Caledonia, Society Is). 

To these must be added: South America (Colombia in 1988 (D. Moore 

pers. comm. 1989), Ecuador (CIBC 1988a, b)); Central America (El 

Salvador, Mexico (Baker 1984)); Asia (India (Kumar et al. 1990)) and the 

Pacific (Fiji (Anon. 1979a), Tahiti (Johnston 1963)). In the West Indies, 

Reid (1983) reported the beetle from Jamaica and Puerto Rico, but it has not 

been reported from the lesser West Indies (Guadeloupe). 

Significant coffee-growing or potential coffee-growing areas not yet 

infested are Solomon Is, Vanuatu, Hawaii and Papua New Guinea, although 

the latter is at serious risk because it shares a common land frontier with Irian 

Jaya (Indonesia), where H. hampei has been present for many years 

(Thomas 1961). H. hampei is not present in Australia. 

The following description of the life cycle refers exclusively to the 

relationship of the beetle with Coffea spp., and principally with Arabian 

coffee C. arabica and robusta coffee C. canephora, 

the most important 

cultivated species. Infestations of H. hampei occur in coffee seeds while 

they are enclosed in berries on the trees and in berries that fall to the ground. 

They will also continue vigorously in processed beans in storage, but not if 

the moisture content has been reduced below 12.5% (robusta beans) or 

13.5% (arabica beans) (Hargreaves 1935). Apart from dispersive flight by 

adult females and the walking by males from one berry to another on the 

same branch (P. Cochereau pers. comm. 1995), no part of the life cycle of 

the coffee berry borer is passed through outside of the coffee bean. 

The length of adult females of H. hampei of American origin is given as 

1.4 to 1.7 mm (Wood 1982) and of Ugandan females and males as about 

1.9 mm and 1.3 mm respectively (Hargreaves 1926). Malaysian females 

averaged 1.58 mm and males 0.99 mm (Corbett 1933). Females outnumber 

males by at least 10 to 1 and the ratio is frequently much higher. The beetle is 

brown when it first emerges from the pupa but in the course of 4 or 5 days it 

becomes generally black, although the prothorax has a slightly reddish tinge. 

The prothorax is markedly humped, so that the down-turned head is not 

visible from above. The tibiae have strong spines which doubtless assist in 

such activities as tunneling through the pulp of coffee berries, ejecting the 

resulting frass, and forcing a way to the soil surface should the berry become 

interred. 

Beetle attack tends to be aggregated on some trees or on particular 

branches within trees, rather than evenly distributed (Baker 1984). The


fertilised female flies to coffee berries that have begun to ripen and bores an 

entrance hole at the apex, either in the terminal pore or in the calyx ridge or 

annulus of differentiated tissue that surrounds the pore. Sometimes this 

annulus is perforated by several holes, but boring into the fruit elsewhere is 

unusual. The colour of berries appears not to influence choice by females 

seeking oviposition sites (Morallo-Rejesus and Baldos 1980). Young 

berries, containing seeds with a watery endosperm, usually do not come 

under attack if more advanced berries are plentiful. If they do they are soon 

abandoned, after the female has fed on some of the pulp, and they then tend 

to fall prematurely, being particularly vulnerable to infection by disease 

organisms. The falling of such immature berries after being attacked often 

contributes significantly to the amount of crop lost. After the endosperm has 

passed from the watery to the milky stage in the course of maturation, beetles 

invading the berry will wait in the pulp until the seed tissue is firm enough to 

excavate (Penatos and Ochoa 1979). Rhodes and Mansingh (1981) cite 

opinions to the effect that females that become static in this fashion for 

several weeks (May to mid-July in the Jamaican lowlands) are in a state of 

reproductive diapause. When available, berries are selected that are already 

suitable for colonisation. The green berry is favoured for feeding and the ripe 

(i.e. red) berry for breeding purposes, but the ripe berries are also very 

suitable for feeding (Corbett 1933). In a ripe berry the female bores in one 

operation through skin, pulp and the endocarp and pellicle surrounding one 

of the two seeds (beans) present in each berry. Ejected frass may surround 

the entrance hole during boring (Hutson 1936). Several days may be 

occupied in this boring process, and the female then tunnels into the 

endosperm, the substance of the seed, which is the basis of the worldÕs US$8 

billion annual coffee crop (Bardner 1978). Berries that fall to the ground 

may generate considerable numbers of beetles, but these are from their ontree 

infestation, since the female berry borers do not appear to visit fallen 

fruit (Baker 1984). 

The eggs are laid at the rate of two or three a day in batches of 8 to 12 in 

chambers chewed out of the maturing bean tissue. Oviposition extends over 

a period of three to seven weeks, each female producing from about 30 to 

over 70 eggs. According to some authors, laying is not necessarily confined 

to one bean because the female that has initiated an infestation may fly to 

other berries during the oviposition period. According to others (e.g. 

Bergamin 1943) the female that has initiated an infestation only quits the 

bean when the first of her progeny emerge as adults. Others again (e.g. 

Hargreaves 1935) state that she remains until all the bean tissue is consumed 

or has deteriorated in some way. Most likely the pattern is quite flexible. 

Eggs hatch in three to nine days and young larvae bore into intact bean

4.10 


161 

tissue, making pockets opening off the main tunnel made by the parent 

female. Male larvae pass through their two instars in the course of about 15 

days, and the females pass through three instars in about 19 days (Bergamin 

1943). Morallo-Rejesus and Baldos (1980) state that the female, like the 

male, passes through only two instars, indicating the need for further 

biological study. The long period over which oviposition is spread results in 

larvae in all stages of development being present in one bean. At the end of 

the larval stage there is a non-feeding or prepupal stage lasting about two 

days. The insect then pupates, without any cocoon formation, in the galleries 

excavated by the larvae. The pupal stage is passed through in four to nine 

days. The period from egg-laying to the emergence of the adult is 25 to 35 

days. The temperatures at which the preceding records were made are 

generally not specified, but chiefly they relate to warm lowland coffee 

plantations. Bergamin (1943) recorded that at 24.5¡C in Brazil the period 

from egg-laying to emergence of adult averaged 27.5 days. De Oliveira 

Filho (1927) found that in Brazil shade temperatures of 20 to 30¡C suited the 

females best. Below 15¡C they became inactive, endeavouring to hide, 

preferably in coffee berries, but sometimes by boring into beans, maize, 

peanuts or cotton seed of suitably low moisture content. They can survive 

temperatures just below 0¡C, which however are rarely experienced in 

Brazilian coffee growing areas. At higher elevations development is 

somewhat prolonged (Le Pelley 1968) and H. hampei has a low pest status in 

highland coffee growing areas in East Africa and Java (Haarer 1962). Baker 

et al. (1989) conclude that the optimum mean annual temperature for the 

beetle is 23¡ to 25¡C and that parasitoids for biological control should be 

sought from a similar climate. 

The adult males emerge from the pupa earlier than the females. Their 

hindwings are short and they do not fly, but remain in the bean, fertilising 

their female siblings as they emerge. Each male can fertilise two females a 

day and up to 30 in his lifetime which may extend to 103 days, although 

averaging less. Corbett (1933) states that the males seldom leave the berries, 

and then only when they are near death. The vast majority of observers 

confirm that males never leave the berries. Quite likely they may move from 

bean to bean within a fruit, thereby gaining access to females other than their 

sisters. Parthenogenesis does not occur and, although unfertilised females 

may produce some eggs, these do not hatch. One insemination is sufficient to 

allow a female to lay fertile eggs throughout her reproductive period. 

Corbett (1933) stated that, if there are no males in the seed when the females 

emerge from the pupal skin after their hardening period of a few days, they 

leave via the entrance hole and seek males in other infested berries. Morallo- 

Rejesus and Baldos (1980) suggest that sex pheromones secreted by the 

males guide such females to appropriate berries.


Females that have been fertilised remain in the ÔparentalÕ bean for three 

or four days, by which time they have become sexually mature. They then 

leave the berries via the entrance holes and enter others and, after a 

preoviposition period of 4 to 20 days, commence egg laying. Females have 

been known to live up to 282 days, and longevity was stated by Bergamin 

(1943) to average 156 days. According to Corbett (1933), in Malaysia 

females survived 81 days without food. There is time for a succession of 

seven or eight generations a year in lowland coffee growing areas but, on 

account of the long reproductive period, there are few clearcut population 

peaks to indicate generations. 

Life history studies have been carried out with artificial infestations of 

coffee trees in southern Mexico (Baker et al. 1992). Morallo-Rejesus and 

Baldos (1980) observed in the Philippines that beetles are to be observed in 

flight from 3.00 pm, considerable numbers being visible in the air between 

4.00 and 5.00 pm. Corbett (1933) observed in Malaysia that females fly at 

any time during the day, but in greatest numbers between 2.00 and 5.00 pm, 

reaching a peak between 3.30 and 4.30 pm. De Oliveira Filho (1927) states 

that, in Brazil, females Ôare activeÕ on warm nights, but it is unclear whether 

this implies flight activity. Kalshoven (1981) states that, in Java, females 

start flying during the midday period, and that they assemble under leaves 

and in other places where they dance up and down like gnats. Such activity 

can have no sexual significance, seeing that the males do not leave the seeds, 

and its function is obscure. In Java flights up to 345 m have been measured 

(Leefmans 1920). In Mexico, Baker (1984) carried out experimental studies 

on flight. Females flew freely in the laboratory for up to 22 minutes, tending 

to hover or move forward only slowly. In tethered flight, and thus relieved of 

supporting their own weight, they could fly non-stop for 100 minutes, with a 

combined aggregate of three hours. Such enduring activity, combined with 

its afternoon peak of activity, suggests that, in their habits, the beetles 

resemble aphids and thrips in being adapted to exploiting periods of 

maximum convection in the atmosphere, so achieving long-distance travel 

with their own contribution serving chiefly to keep them aloft. De Oliveira 

Filho (1927) states that local flight occurs when the fertilised female is 

seeking a place to lay, when (oviposition having commenced) she emerges 

to seek moister berries after having been driven out by the heat of the sun. It 

also occurs when unfertilised females seek males (as they do if there are 

none in the berry when they emerge), when seeds are waterlogged, are 

overcrowded with adults and larvae, or when the beetles are disturbed. 

Rhodes and Mansingh (1981) state that, in the Jamaican lowlands, 

beetles in dry berries remain in diapause for five months, from mid- 

December to mid-May. Baker (1984) found that in mid-spring in Mexico

Host plants 

4.10 


163 

females tended to remain in fallen coffee berries at a time when temperatures 

in berries in the trees ranged up to an inimical 37¡C. Soaking the fallen 

berries in water induced many to emerge, but they did so in a specific 

pattern, some seven to eight hours after dawn. Possibly the soaking 

simulated rain that would have made the environment generally more 

favourable. Baker reminds us that coffee is naturally an understorey plant in 

tropical forest and, by sheltering in fallen berries, beetles may avoid the 

harmful effects of strong, direct sunlight. Infestations are carried over 

between peaks of fruiting by the breeding that occurs in late-maturing 

berries, or else in those that have fallen to the ground. Females can survive 

for up to two months in buried beans (Clausen 1978). 

It is probable that intercontinental travel is brought about by the agency 

of man, rather than by travel in moving air masses. Infested beans are an 

obvious vehicle for dispersal, but there are other avenues to which 

quarantine measures should be applied. In Jamaica, Reid (1983) observed 

females among banana trash used in packing boxes on their way to the 

boxing plant. Commonly, beetles disperse in sacks, empty or otherwise, and 

on the clothing and equipment of plantation workers. Under some conditions 

beetles bore for protection into wood or other materials to the extent that 

Baker (1984) suggested that authorities in beetle-free areas should think 

very carefully before allowing entry of untreated plant material from an 

infested area. 

An important aspect of the biology of any insect pest is its host range. In 

Africa, in addition to its regular hosts in the genus Coffea, 

Hypothenemus 

hampei has been reported from fruit, pods or seeds of species of Centrosema, 

Crotalaria, 

Phaseolus and Tephrosia (Fabaceae), Leucaena (Mimosaceae), 

Caesalpinia (Caesalpiniaceae), Hibiscus (Malvaceae), Rubus and 

Oxyanthus (Rubiaceae), Vitis (Vitaceae) and Ligustrum (Oleaceae), but 

these associations are all considered to reflect only casual feeding by adults. 

In Africa, the only species outside of the genus Coffea in which immature 

stages have been found is Dialium lacourtianum (Caesalpiniaceae) (Le 

Pelley 1968). 

A review of hosts of the genus Hypothenemus was made by Johanneson 

and Mansingh (1984) who concluded that H. hampei was monophagous 

according to their criteria, as it attacked only six species of the genus Coffea. 

However, they listed 23 other species of plants in 11 families from which 

H. hampei has been recorded, but only as adult females. In contrast, in the 

Philippines, Morallo-Rejesus and Baldos (1980), whose paper was


Damage 

overlooked by Johanneson and Mansingh, reported finding eggs, larvae and 

pupae of H. hampei in Leucaena leucocephala (Mimosaceae), Gliricidia 

sepium (Fabaceae), two species of Psychotria (Rubiaceae) and one of 

Dioscorea (Dioscoreaceae). In laboratory tests they found that adults of 

H. hampei fed on pods of four of those species and also on the pods of 19 

other species in 9 orders. 

Such feeding tests may be of little significance, however, since the 

survival times recorded are greatly exceeded by the periods for which the 

beetles are capable of withstanding starvation (Corbett 1933). If the insects 

were correctly identified, the host plants recorded in the Philippines may 

help to support a population of H. hampei when no coffee berries are 

available. Reexamination of the host range is necessary. For example Cohic 

(1958) found H. hampei attacking loquat in New Caledonia, and this 

relationship, though abortive in the end, has not been reported anywhere else 

in the world. In connection with host records, Johanneson and Mansingh 

(1984) drew attention to the problem of misidentification of species of 

Hypothenemus, 

a notoriously difficult genus, and also to misinterpretation 

of the relative roles of various host plants. Hargreaves (1935) found adults of 

four species of Hypothenemus other than H. hampei in seed of Phaseolus 

lunatus (Fabaceae) in Uganda, and Gonzalez (1978) alludes to species of 

Hypothenemus, 

known as false coffee borers, which occur from Mexico to 

northern Argentina and greatly complicate quarantine procedures. Such 

insects would, of course, also raise difficulties in host plant studies. A 

thorough review of true hosts of H. hampei would be relevant to a number of 

aspects of the control of this pest. 

Hypothenemus hampei is a pest exclusively of the immature and mature 

coffee berries and does no damage whatsoever to the vegetative parts of the 

plant. Prates (1969) showed that adults of H. hampei were strongly attracted 

to extracts of green or ripe coffee berries, but not to extracts of coffee leaves 

or flowers. Significant losses are caused by the female beetles feeding on 

young berries which are too immature to colonise but which, after the beetle 

has gone, are invaded by decay organisms, and so fall prematurely. In Java 

Leefmans (1920) found that 80% of green berries that had fallen through 

being bored by the beetle contained decayed beans as against 46.5% in 

unbored beans that had fallen through other causes. In the Congo, Schmitz 

and Crisinel (1957) found that 64 to 82% of shed berries had fallen on 

account of H. hampei attack. Such losses caused by attack on immature 

fruits are serious enough, but the bulk of the damage done by this beetle is to

4.10 


165 

the endosperm of the mature beans, which may be extensively damaged or 

even completely destroyed. Even lightly bored beans acquire a distinctive 

blue-green staining which significantly reduces their market value (McNutt 

1975), but the further tunnelling by the beetles and their larvae brings about 

progressive degradation, so that the coffee bean is reduced to a mass of frass. 

Market requirements demand the removal of damaged berries from the 

harvested crop, which is done by various mechanical processes (fortunately 

bored beans float), supplemented even by handpicking. The beans removed 

by such processing are not necessarily a total loss, but can go into only low 

grade fractions at a much reduced market rate. 

In New Caledonia, where no control measures had been implemented, 

H. hampei was found to have attacked 80% of berries (Cohic 1958). Other 

examples of losses due to Hypothenemus hampei are given by Le Pelley 

(1968). Severe infestations in Uganda may result in 80% of berries being 

attacked. In the Ivory Coast, damage of 5% to 20% of berries is common, 

rising to 50% to 80% in some cases. In the Congo, boring of up to 84% of 

green berries and up to 96% of hard berries has been recorded and, in 

Tanzania, records indicate up to 96% boring of hard berries. In Malaysia 

there have been records of up to 90% of beans damaged. In Java crop loss of 

40% was recorded in 1929, and in Brazil 60% to 80% losses have been 

experienced. The above figures apply for the most part to poorly managed 

situations, and crop losses can be reduced by appropriate management, but 

the beetle is a constant latent threat if vigilance is relaxed. In Jamaica, Reid 

(1983) estimated that 27% of the berries harvested were damaged. The 

studies of Reid and Mansingh (1985) showed that H. hampei was 

responsible for 20.9% reduction of exportable beans in the Jamaican crop of 

1980Ð81. Baker (1984) reported that, in southern Mexico, the attack of 

H. hampei on coffee plantations was so severe that, in spite of application of 

insecticides in some places in 1982, no berries were harvested because it 

would not have been economical to do so. 

Proper processing results in beans of moisture content too low to permit 

the borer to multiply. This is below 13.5% for arabica coffee and below 

12.5% for robusta coffee. If coffee beans are stored with significantly higher 

moisture content, beetle reproduction continues. Thus Morallo-Rejesus and 

Baldos (1980) found that, in the Philippines, infestation in coffee beans 

stored before drying rose from 20% to 100% in six weeks.



The cryptic nature of the immature stages and the male of H. hampei makes 

them relatively inaccessible victims for predators, and the only one recorded 

is the non-specific Javanese bug Dindymus rubiginosus. This bug draws the 

borers from the berries with its beak and sucks them dry. Le Pelley (1968) 

states that it is of little importance. 

The most important parasitic wasps, Cephalonomia stephanoderis, 

Prorops nasuta, Phymastichus coffea and Heterospilus coffeicola are, of 

course, African in origin and are dealt with in some detail by Klein Koch et 

al. (1988) and Feldhege (1992). C. stephanoderis which is restricted to West 

Africa is the most important species in Ivory Coast, parasitising up to 50% of 

H. hampei in black berries (Ticheler 1961). The potential of H. coffeicola in 

biological control requires further study because its larvae are not very 

specific, but the other three species appear to have a narrow enough host 

range to make them acceptable from this point of view. A fifth parasite, 

Goniozus sp. is recorded, but without further data, from Ivory Coast 

(Cochereau and Potiaroa 1994). 

In addition to the identified arthropod natural enemies (Table 4.10.1), 

Leefmans (1924a) recorded a non-specific parasite that attacks beetles in 

newly infested berries and Hargreaves (1926) found an unidentified 

hymenopterous parasitoid in Uganda, now known also from Togo as 

Aphanogmus dictynna and considered to be a hyperparasitoid of 

C. stephanoderis or P. nasuta (Feldhege 1992). Morallo-Rejesus and 

Baldos (1980) reported the presence in the Philippines of a braconid and an 

encyrtid parasitoid of H. hampei, both unidentified, and presumably nonspecific 

members of the local fauna. 

Some ants attack the borer. Swallows and other small birds that feed on 

the wing consume flying adults of H. hampei. 

The parasitic fungus Beauveria bassiana has been observed attacking 

H. hampei in Brazil (Averna-Sacc‡ 1930; Villacorta 1984), Jamaica 

(Rhodes and Mansingh 1981), Cameroon (Pascalet 1939), Congo (Sladden 

1934; Steyaert 1935), Ivory Coast (Ticheler 1961), India (Balakrishnan et al. 

1994), Java (Friederichs and Bally 1922) and in New Caledonia (Cochereau 

and Potiaroa 1994). Steyaert (1935) and Averna-Sacc‡ (1930) studied the 

seasonal cycle and the former also made studies of the infectivity and 

epidemiology of the fungus of which there are many strains. In an analysis of 

16 isolates from H. hampei adults from 10 countries in Latin America, 

Africa, Asia and the Pacific, 13 formed a homogenous group with very 

similar electrophoretic and physiological characteristics, suggesting a 

distinct strain associated widely with the coffee berry borer. Of the

4.10 Hypothenemus hampei 167 

remaining 3 strains, one from Sri Lanka is suspected as having degenerated 

during some 63 years in storage, but the others (from New Caledonia and 

Kenya) are probably distinct entities (Bridge et al. 1990). The New 

Caledonian strain presumably attacked some other host until H. hampei 

arrived there in 1948. It is a particularly virulent strain and can cause death of 

H. hampei in 5 days (Cochereau et al. 1994). Moist, warm conditions favour 

the incidence of this pathogen, and heavy rain is thought to enhance the rate 

of infection. If spraying with fungal preparations is avoided on the day of 

release of parasitoids, adverse effects on the latter are not observed (Reyes et 

al. 1995). Friederichs (1922) recommended the encouragement of heavy 

shade to increase the incidence of fungal pathogens, but this runs counter to 

the fact that intensity of shade must often be reduced to encourage 

hymenopterous parasitoids which, however, may still prove to be of minor 

significance in population regulation. Certainly, Klein Koch (1989a) 

considered Beauveria to be the most important natural enemy of H. hampei 

in Ecuador. In Colombia, preparations of selected strains of Beauveria in oil 

have produced 20 to 95% adult mortality, slightly higher than the 20 to 90% 

produced by selected strains of Metarhizium (P. Cochereau, pers. comm. 

1995). Varela and Morales (1996) have characterised a number of Beauveria 

isolates and their virulence against H. hampei. 

Pascalet (1939) advocated the spraying of suspensions of spores, before 

sunrise, but no results are available. As with so many parasitic fungi, its 

application would be limited by intolerance of dry conditions. Another 

fungus that attacks H. hampei, Paecilomyces javanicus, is Afro-Asian in 

distribution and wide spectrum in its host range (Samson 1974), attacking 

also Lepidoptera. Its use against H. hampei appears not to have been 

attempted. 

There appears to be only one record of nematodes attacking H. hampei in 

the field (Varaprasad et al. 1994), but in addition, Allard and Moore (1989) 

showed that a Heterorhabditis sp. could cause high mortality of both adult 

and larval H. hampei under laboratory conditions and that infective 

juveniles were produced from adults and larger larvae. Spraying of 

nematodes on fallen berries might remove the need to collect them (which 

involves much labor), leaving them to provide mulch. Dispersal of infected 

adults may also spread the nematodes into the pest population. Further work 

with nematodes is clearly desirable.

Table 4.10.1 Natural enemies of Hypothenemus hampei 

Species and type 

HEMIPTERA 

Country Reference Comment 

PYRRHOCORIDAE (Predator) 

Dindymus rubiginosus 

HYMENOPTERA 

Java Wurth 1922 Not specific 

BETHYLIDAE (ectoparasites of immature stages) 

Cephalonomia stephanoderis Ivory Coast Betrem 1961; Ticheler 1961; 

Cochereau & Potiaroa 1994; 

A promising parasite 

Togo 

Klein Koch et al. 1988 

Goniozus sp. Ivory Coast Cochereau & Potiaroa 1994 

Prorops nasuta Cameroon Klein Koch et al. 1988 

Congo 


Ivory Coast Klein Koch et al. 1988 

Kenya 


Tanzania Rangi et al. 1988 

Togo 


Uganda 

Klein Koch et al. 1988; 

Klein Koch et al. 1988; 

Waterston 1923 

Scleroderma cadaverica 

CERAPHRONIDAE 

Uganda Benoit 1957 Causes severe dermatitis in man 

Aphanogmus (= Calliceras) dictynna 

EULOPHIDAE 

Uganda Waterston 1923 Possibly hyperparasitic 

Phymastichus coffea (attacks 

Ivory Coast Cochereau & Potiaroa 1994 

adult beetles) 

Kenya 

La Salle 1990 

Togo 



Table 4.10.1 (contÕd) Natural enemies of Hypothenemus hampei 

Species and type 

HYMENOPTERA 

Country Reference Comment 

BRACONIDAE 

(ectoparasitoid and predator) 

Heterospilus coffeicola 

Uganda 

Tanzania 

Cameroon 

Congo 

Schmiedeknecht 1924 

CIBC 1988b 



Kills larvae with sting 

Attacks larvae of other parasites of 

H. hampei Ñ also may be 

cannibalistic 

FORMICIDAE 

(predator) 

Crematogaster curvispinosa 

ACARI 

Brazil Pinto da Fonseca & Araujo 1939 Can cause high mortality of immature 

stages in coffee berries 

Pyemotid mite 

NEMATODA 

New Caledonia P. Cochereau pers. comm. 

Heterorhabditis sp. Moore & Prior 1988 

Panagrolaimus sp. 

FUNGI 

India Varaprasad et al. 1994 

HYPHOMYCETES 

Beauveria bassiana 

(= Botrytis stephanoderis) 

Java 

Cameroon 

Friederichs & Bally 1922 

Pascalet 1939 

Cosmopolitan, in a variety of strains 

Metarhizium anisopliae Moore & Prior 1988 

Nomuraea rileyi Moore & Prior 1988 Usually recorded from Lepidoptera 

Paecilomyces (= Spicaria) javanicus Java Friederichs and Bally 1922; 

Samson 1974 

Indonesia, Asia, Africa 

P. tenuipes Moore & Prior 1988 

4.10 Hypothenemus hampei 169



Africa 

Published information is summarised in Table 4.10.2, but there were 

probably a number of transfers of parasites within Africa and perhaps South 

America that have gone unrecorded. In the past decade the International 

Institute for Biological Control had adopted the policy of breeding African 

parasitoids in England on H. hampei in coffee beans from the country of 

destination. This is because of the possibility that the wasps might carry 

spores of fungal diseases of coffee, especially new strains of coffee leaf rust 

(Hemileia vastatrix) and coffee berry disease (Colletotrichium coffeanum) 

(Moore and Prior 1988; Rangi et al. 1988; Nemeye et al. 1990; Murphy and 

Rangi 1991). The danger of fungal transmission could also be reduced by 

breeding H. hampei on an artificial diet (Brun et al. 1993; Perez et al. 1995; 

Villacorta 1985). 

CAMEROON 

Pascalet (1939) recommended the introduction of Heterospilus coffeicola, 

Prorops nasuta and Beauveria bassiana to any plantations lacking them. 

There is no record that this was implemented anywhere, nor whether any or 

all of the organisms were not already generally present. 

CONGO 

Sladden (1934) and Leroy (1936) suggested that, by breeding and liberating 

them, it would be possible to increase the efficiency of P. nasuta and 

H. coffeicola, which he knew to be already present in the Congo and he made 

a similar suggestion for fungus diseases. However, there is no indication of 

the extent to which this was done. 

KENYA 

Prorops nasuta was sent from Uganda to Kenya in 1930 (Greathead 1971), 

but according to Evans (1965) that wasp and H. coffeicola were probably 

native there. Abasa (1975) considered that parasites were of doubtful value 

in controlling H. hampei in Kenya. 

UGANDA 

Prorops nasuta and Heterospilus coffeicola are both native to Uganda. 

Hargreaves (1935) considered that some areas lacked these parasites, and so 

he introduced cultures from Kampala County, north of Lake Victoria, to 

Bwamba County on the western border. He stated that this introduction 

resulted in a great reduction in the previously intense infestation of coffee 

berry borer but, in view of the natural occurrence of P. nasuta over a wide 

area to the west of the Ugandan border (Le Pelley 1968), it seems unlikely 

that the distribution was discontinuous and that it was lacking in Bwamba

Asia 


County. HargreavesÕ claims that the introduction brought about a great 

reduction in the impact of the coffee berry borer in Bwamba County must be 

treated with reserve, the more so since De Toledo Piza and Pinto da Fonseca 

(1935) state that neither P. nasuta nor H. coffeicola appeared to control the 

borer in nearby Kampala. More recently, P. nasuta was reported to achieve 

20% parasitisation in western Kenya in the dry season (Barrera et al. 1990b). 

SRI LANKA 

Stock of P. nasuta and H. coffeicola from Uganda were liberated in Sri 

Lanka in 1938, but neither species became established (Hutson 1939). 

INDONESIA 

The introduction of Prorops nasuta to Java from Uganda in 1923 was the 

earliest attempt to bring about the biological control of H. hampei which had 

first been reported in Java in 1909 (Kalshoven 1981). P. nasuta was found to 

be easily propagated (Leefmans 1924a), was distributed widely in 

considerable numbers (Begemann 1926) and became established (Le Pelley 

1968). However, it apparently could not maintain itself and was still being 

bred for distribution in 1928 (UltŽe 1928) and in 1932 (Betrem 1932; 

Schweizer 1932; UltŽe 1932). 

Leefmans (1924a) drew attention to the fact that the P. nasuta did not 

thrive in shade, and that it flourished best in black berries which tend to be 

most abundant after harvest, when the parasite is least needed. The former 

problem was solved by appropriate pruning but, despite improvements in 

management to favour it and the long period spent in breeding and 

disseminating it, the parasite seems not to have become established in Java 

(Clausen 1978; Kalshoven 1981). 

Cultures of Heterospilus coffeicola were taken to Java from Uganda 

along with those of P. nasuta in 1923, but the wasp appears not to have been 

released. Leefmans (1924a) seems to have concluded that it was likely to be 

incompatible with P. nasuta.

Table 4.10.2 Introductions for the biological control of Hypothenemus hampei 

Country and species liberated 

BRAZIL 

Year From Result Reference 

Cephalonomia stephanoderis ? ? + Benassi & Berti-Filho 1989 

Prorops nasuta 

COLOMBIA 

1929 Uganda + Hempel 1933; Yokoyama et al. 1978 

Cephalonomia stephanoderis 1988 Kenya via U.K. + C. Klein Koch pers. comm. 1995; 

Sponagel 1993 


ECUADOR 

1995 + Bustillo et al. 1995; 

Portilla & Bustillo 1995 

Cephalonomia stephanoderis 1988 Togo via U.K. + CIBC 1988b; Klein Koch 1989a,b,c; 

Klein Koch et al. 1988; Delgado et al. 

1990; Sponagel 1993 

Prorops nasuta 1987Ð1990 

1988 

Kenya via U.K. 

West Africa 

Ð 

Ð 

CIBC 1988a; Klein Koch et al. 1988; 

Rangi et al. 1988; Murphy & Rangi 

1991; Sponagel 1993 

EL SALVADOR 

Cephalonomia stephanoderis 

GUATEMALA 

1988 Kenya via U.K. + Sponagel 1993; C. Klein Koch pers. 

comm. 1995 

Cephalonomia stephanoderis 1988 

1993-1995 

HONDURAS 

Kenya via U.K. ? Sponagel 1993 

Garcia & Barrios 1996 

Cephalonomia stephanoderis 1988 Kenya via U.K. ? Sponagel 1993 


Table 4.10.2 (contÕd) Introductions for the biological control of Hypothenemus hampei 

Country and species liberated 

INDONESIA 

Year From Result Reference 

Cephalonomia stephanoderis 1988 ? Sponagel 1993 

Heterospilus coffeicola 1923 

1931 

Prorops nasuta 1923Ð 

1925 

KENYA 

Uganda 

Uganda 

not liberated 

? 


Le Pelley 1968 

Uganda Ð Begemann 1926 


Prorops nasuta 1930 Uganda already present Evans 1965 

MEXICO 

Cephalonomia stephanoderis 1988Ð1989 Togo via U.K. + Barrera et al. 1990 a, b; 

CIBC 1988b 


NEW CALEDONIA 

1988Ð1989 Kenya and Togo via U.K. ? Barrera et al. 1990a, b; 

Murphy & Rangi 1991 

Cephalonomia stephanoderis 

UGANDA (BWAMBA COUNTY) 

1993 Ivory Coast + Cochereau & Potiaroa 1994 


PERU 

1932 Uganda (Kampala county) + 

?already present 

Prorops nasuta 1962 

1964? Brazil Ð 

SRI LANKA 

Hargreaves 1935 

Clausen 1978 

De Ingunza 1964 

Heterospilus coffeicola 1938 Uganda Ð Hutson 1939 

Prorops nasuta 1938 Uganda Ð Hutson 1939 

4.10 Hypothenemus hampei 173


Pacific 

NEW CALEDONIA 

Infestation of coffee berries with H. hampei ranges from 0% to 100%, with 

an overall average of 33%. C. stephanoderis from West Africa was released 

in 1993 and recovered almost a year later. However, wherever the 

aggressive, little red fire ant (Wasmannia auropunctata, introduced around 

1970) is present this parasitoid is unable to survive. When the fire ant is 

eliminated from a plantation by banding the trees with insecticide the wasp is 

established. Ant control is, thus, a prerequisite for biological control 

(Cochereau and Potiara 1994; P. Cochereau pers. comm. 1995). 

Phymastichus coffea is currently under consideration for liberation 

(P. Cochereau pers. comm.). Cochereau et al. (1994) have examined in some 

detail the effectiveness against H. hampei of a virulent New Caledonian 

strain of Beauveria bassiana, which shows considerable promise in the field. 

Central America 

GUATEMALA 

Cephalonomia stephanoderis was mass produced and released during 1993 

to 1995. Infestation by H. hampei was reduced 75%, to 2.7 to 5.3% in 1993 

and to 0.4 to 0.9% in 1994, but an increase was observed in 1995 (1.6 to 

2.4%, compared with the control of 3.8% infestation), resulting in 48% 

control (Garcia and Barrios 1996). The cost of mass liberations of 

C. stephanoderis was comparable with that of chemical control (Decazy et 

al. 1995). 

MEXICO 

Cephalonomia stephanoderis from Togo and Prorops nasuta from both 

Togo and Kenya were raised in U.K. in coffee beans from Mexico before 

being sent there for mass production and liberation during 1988 and 1989. 

C. stephanoderis has become established, but the situation with P. nasuta is 

unclear (Barrera et al. 1990a, b). The impact of these parasitoids remains to 

be reported. 

South America 

BRAZIL 

Prorops nasuta was imported into Brazil from Uganda in 1929, and by 1933 

it was established in several coffee plantations (Hempel 1933, 1934). As in 

Java, breeding and distribution continued and in 1937 (Anon. 1937) it was 

stated to be of considerable value in controlling the coffee berry borer in S‹o 

Paulo, but only if its numbers were boosted by rearing between coffee 

production seasons. Puzzi (1939) studied the reproduction of the parasite in


relation to that of its host in Brazil and concluded that, in theory, it was more 

prolific, but that the efficiency of the parasite was limited by the tendency of 

the female to remain in one berry. De Toledo (1942) examined rates of 

parasitisation, but his figures do not suggest that the wasp could have been 

having any significant impact. De Toledo et al. (1947) were only mildly 

enthusiastic about the value of the wasp, mentioning a continuing need for 

repeated liberations and the requirement for boosting the effect of the 

parasite by cultural practices. Le Pelley (1968) stated that, at that time, 

Brazilian entomologists appeared satisfied that P. nasuta was of value in 

their country, but he could find no conclusive evidence that the amount of 

routine work required for the control of H. hampei had decreased. 

Yokayama et al. (1978), considered that the climate of the S‹o Paulo district 

in Brazil was unfavourable for this wasp, in which the growers lost interest 

when BHC was found to give satisfactory control. Nevertheless, they 

reported that it had recently been recovered in coffee plantations in S‹o 

Paulo, having survived pesticide usage, severe droughts and winter frosts. 

The fact that P. nasuta has been transferred more recently to areas in Brazil 

where it is not established indicates that the wasp is considered to be of some 

value (Ferreira and Batistela Sobrinho 1987). 

Although there is no record of its release, a species of Cephalonomia, 

presumably C. stephanoderis was recovered in the field in Brazil between 

1986 and 1988 (Benassi and Berti-Filho 1989). 

A survey of natural enemies of H. hampei in northern Espirito Santo 

from 1986 to 1994 revealed 3 parasitoids (Prorops nasuta, Cephalonomia 

sp., and a species of Proctotrupoidea), an ant predator (Crematogaster 

curvispinosa) and a fungus (Beauveria bassiana) (Benassi 1995). 

COLOMBIA 

The rate of population increase of H. hampei in the field has been studied by 

Gaviria et al. (1995). A major program of integrated management 

commenced in 1992, involving local strains of the fungi Beauveria bassiana 

and Metarhizium anisopliae and parasitoids, in particular Cephalonomia 

stephanoderis and Prorops nasuta (CABI: IIBC 1993; C. Klein Koch pers. 

comm. 1994; Bustillo et al. 1995). Methods for mass production of 

C. stephanoderis and Prorops nasuta are provided by Portilla and Bustillo 

(1995). Sixty million parasitoids were released and 100 tons of B. bassiana 

and M. anisopliae were applied (Bustillo et al. 1995). 

ECUADOR 

Klein Koch (1986) proposed the introduction of 3 parasitoids from Africa, 

Prorops nasuta, Heterospilus coffeicola and Cephalonomia stephanoderis. 

C. stephanoderis was first introduced from Togo in 1988 and P. nasuta from 

Tanzania and Kenya in 1987Ð88 and Togo in 1988 (Klein Koch 1989c).


PERU 

Liberations continued in subsequent years with some 920000 of the former 

species and 30,000 of the latter being released in 1992. Both species are now 

well established and having a significant effect. In field experiments with 

caged coffee trees 86% of berries on the bush and 87% on the ground 

contained C. stephanoderis and up to 52% and 31% parasitisation 

respectively was recorded from tree and ground berries in the open (Klein 

Koch 1989b,c, 1990). However the results are poor in the Amazon region of 

the country where the rainfall is very high. Elsewhere, as part of an 

integrated approach, infestation levels on Coffea arabica are as low as 0.4 to 

1.4%. Recently introduced catimor varieties are resistant to coffee rust, so 

sprays are no longer required (Klein Koch pers. comm. 1994). In addition, 

the fungus Beauveria bassiana and the ant Azteca sp. cause considerable 

mortality of H. hampei when humid weather conditions prevail (Klein Koch 

et al. 1987). 

According to De Ingunza (1964) Prorops nasuta was introduced from Brazil 

to Peru in 1962, but failed to become established. 

Major parasite species 

Cephalonomia stephanoderis Hym.: Bethylidae 

Cephalonomia stephanoderis is a small black bethylid wasp which is native 

to West Africa (Ivory Coast, Togo). The females, which are 1.6 to 2.0 mm in 

length, enter bored coffee berries and deposit eggs on the ventral surface of 

final stage larvae and prepupae of H. hampei. Its larvae feed as ectoparasites, 

exhausting the tissues of the host in 4 to 6 days, then spinning a silken 

cocoon in which to pupate. The pupal stage lasts about 15 days. Fertilisation 

takes place in the berry where the wasps emerge, and seemingly, the males, 

although fully winged, remain there after the females have left. Females 

must feed for two days at 27¡C or 6 to 11 days at 24¡C before they can 

mature eggs. Adult females feed by preference on H. hampei eggs and 

young larvae but also on prepupae and they chew holes in the intersegmental 

membrane of adult beetles, between the prothoracic and mesothoracic 

tergites, and feed on the haemolymph. Females cannot produce eggs on a 

diet of borer eggs or adults alone, but need to feed first on the larvae and/or 

prepupae of the borer. They can lay up to 70 eggs (Barrera et al. 1989, 1993; 

Abraham et al. 1990; Wegbe 1990; Infante et al. 1994a ) and can distinguish 

between parasitised and unparasitised hosts (Barrera et al. 1994). In the 

Ivory Coast, Koch (1973) found that adult C. stephanoderis each required 

two eggs, two larvae or two adults per day for survival. At the end of the 

coffee season H. hampei populations were reduced by parasitisation by 20%


to 30%, but by not more than 5% between seasons. Ticheler (1961) recorded 

up to 50% parasitisation by this, the most important parasitoid in the Ivory 

Coast. In West Africa C. stephanoderis is commoner than P. nasuta 

(Abraham et al. 1990). A major mass production and release program 

commenced in Colombia in 1993, with a production of 10 million wasps per 

month. When 400 000 wasps were released in a 2 million ha area of coffee 

trees 85% parasitisation was attained when there was 80% infestation of 

coffee berries and 20% parasitisation when there was 5% infestation. The 

target for the releases was 12.5 wasps per berry and 300 berries containing 

wasps for each 15 trees, giving 20 000 to 30 000 wasps per ha (P. Cochereau 

pers. comm. 1995). Life tables were developed for C. stephanoderis in 

Mexico (Infante and Luis 1993; Infante et al. 1994b). 

Studies of mass releases of C. stephanoderis suggest that they can 

control low density populations of H. hampei in commercial coffee 

plantations, adult predation by the wasp probably being the most important 

mortality factor. However, mass production costs are too high for releases to 

be economically viable and cheap artificial diets for mass rearing are being 

investigated, with successful rearing already achieved for four generations 

(CABI:IIBC 1996). 

Heterospilus coffeicola Hym.: Braconidae 

Heterospilus coffeicola is a braconid wasp about 2.5 mm long. It does not 

enter the borehole of the beetle, but travels from berry to berry inserting its 

ovipositor into the boreholes in the course of seeking Hypothenemus larvae. 

Only one small egg is laid in each berry, and the larva that emerges after 

about six days feeds on beetle eggs and larvae over a period of 18 to 20 days, 

consuming 10 to 15 eggs and larvae per day. In this regard it is more of a 

predator than a parasite. According to De Toledo Piza and Pinto da Fonseca 

(1935) the larva kills the adult H. hampei before pupating inside a white 

silken cocoon. The wasp emerges after a comparatively brief pupal period 

(Hargreaves 1926; De Toledo Piza and Pinto da Fonseca 1935, Le Pelley 

1968). In Uganda it is attacked by a chalcidid of the genus Closterocerus 

(Schmiedeknecht 1924). 

Hargreaves (1926, 1935) stated that Heterospilus coffeicola contributed 

substantially to the control of H. hampei in Uganda. The Brazilian 

entomologists De Toledo Piza and Pinto da Fonseca (1935) studied the wasp 

in Uganda with a view to assessing its potential value as a biological control 

agent in Brazil. They concluded that H. coffeicola can thrive only in areas 

with a continuous production of coffee berries throughout the year, and as 

such conditions prevail nowhere in Brazil they recommended against its 

importation. One possible disadvantage of this wasp is that its larvae feed on


the larvae of other wasps as well as those of H. hampei, and it may even be 

cannibalistic (Hargreaves 1924). If these statements are verified then there 

may be reservations about the employment of H. coffeicola in biological 

control. A further difficulty associated with this species as a biological 

control agent is that a number of workers have been unable to breed it in the 

laboratory, a problem also experienced by CIBC (1987) during its current 

program, although Rangi et al. (1988) have reported limited success. The 

free-living existence of the adults may involve special nutritional or mating 

requirements that have not yet been met experimentally. 

Phymastichus coffea Hym.: Eulophidae 

This parasitoid was first recorded in Togo as recently as 1989 (Borbon- 

Martinez 1989), causing up to 30% parasitisation, but is now known also 

from Ivory Coast and Kenya. It is an endoparasite of the adult female 

H. hampei, which is usually attacked as she is commencing to tunnel in to a 

coffee berry. P. coffea also enters the berry to parasitise male H. hampei. 

Oviposition occurs in both the thorax and the abdomen of the host, the 

former producing a male and the latter a female. Although several eggs may 

be laid in each host, only two wasps are produced. Males range in length 

from 0.45 to 0.55 mm and females from 0.8 to 1.0 mm. Females do not 

require to be fertilised before they commence oviposition shortly after 

emergence. At 27¡C, larval development takes about 21 days, the pupal 

stage about 8 days and adult longevity appears to be a few days only. In the 

field 20 females were produced for each male and it was estimated that 

between 4 and 7 hosts could be parasitised in a 4-hour period. 

P. coffea was the most important parasitoid of H. hampei in the majority 

of coffee holdings on the Togolese Plateau at about 800m above sea level 

and is fairly common around Man (West Ivory Coast) near the Liberian 

border (La Salle 1990; Feldhege 1992; P. Cochereau pers. comm. 1994; 

Infante et al. 1994a, ). 

Mass production methods have been developed in Colombia for 

P. coffea and a decision to release is expected shortly (CABI:IIBC 1996). 

Prorops nasuta Hym.: Bethylidae 

Prorops nasuta is native to Uganda, Kenya, Tanzania and Cameroon, Ivory 

Coast and Togo. It is a dark brown bethylid wasp about 2.3mm in length, the 

name nasuta referring to the characteristic bilobed ÔnoseÕ protruding 

forwards above the antennal bases. This wasp parasitises and preys upon 

several species of Hypothenemus (Clausen 1978). Males that emerge first 

from pupae always stay on the remaining cocoons within the coffee berry 

and mate with the females as they emerge. Because P. nasuta populations 

are, thus, highly inbred it is probable that there are different strains of


P. nasuta in West Africa and East Africa, (Abraham et al. 1990; Griffiths 

and Godfray 1988; Murphy and Rangi 1991) and testing these may be highly 

relevant to biological control. On coffee the fertilised female enters an 

infested berry via the borehole of the adult H. hampei, choosing berries on 

the trees rather than those on the ground. If the parent borer beetle is still 

present she may kill it and use the cadaver to plug the entrance hole, over 

which she stands guard. According to CIBC (1987) the female wasp does not 

feed upon borer beetles she may kill, but other authors state that she will do 

so if no other life history stages are available, but that she cannot mature eggs 

on a diet of adults alone. Several larvae and pupae may be injured with the 

ovipositor before any oviposition occurs, and these victims succumb in a 

few days. P. nasuta feeds by preference on the eggs and young larvae of 

H. hampei and oviposit on the late third stage larvae and pupae. The hosts 

chosen for oviposition are stung, sometimes several times, and thus 

paralysed before one, or sometimes two, eggs are laid upon them. Eggs are 

placed ventrally on larvae and on the abdominal dorsum of pupae. The eggs 

(0.55 ´ 0.18mm) are large for a wasp of this size. They hatch in an average of 

about three days and the larval stages last three to eight days. The 

ectoparasitic larva may consume more than one host. There is a prepupal 

(non-feeding) period of about three days, passed inside a silken cocoon spun 

by the fully fed larva. It is common to find 20 cocoons in a coffee bean, and 

up to 62 have been recorded. The pupal stage lasts on an average about 21 

days, varying from 9 to 27 days according to temperature. 

The life cycle from egg to adult lasts 17 to 33 days (average 29) at 25¡C 

and may be as long as 66 days at 18¡C. There are considerable discrepancies 

between figures given by various authors for the duration of the life history 

stages, but there is general agreement that the female is quite long-lived Ñ 

up to 135 days being cited in Brazil, given an abundant supply of larvae and 

prepupae as food. By contrast it appears that the males do not feed and they 

do not survive longer than 13 days. Females outnumber the males, a figure of 

three to one being recorded. Statements as to duration of the preoviposition 

period give rather diverse figures. Usually a few days are indicated but one 

record is of 17 days. Parthenogenesis may occur, when only male progeny 

are produced. Females may lay up to 66 eggs at a rate of one or two a day, 

utilising several berries in the process. 

In feeding, females consume several eggs and unparasitised larvae per 

day and they will also eat pupae. Normally all stages of the beetle in a berry 

are killed either by parasitisation, predation or merely by stabbing before the 

female leaves (Leefmans 1924a; Begemann 1926; Hempel 1933; De Toledo 

Piza and da Fonseca 1935; Hargreaves 1935; Hutson 1936; Puzzi 1939; De 

Toledo 1942; Le Pelley 1968; Abraham et al. 1990; ). The low abundance of


P. nasuta in Western Kenya suggests that it may not be suited to high 

altitudes (Murphy and Rangi 1991). In one locality in S‹o Paulo (Brazil) the 

percentage of infested berries that contained parasitoids rose to a maximum 

of 2.4 in autumn (De Toledo 1942). 

Scleroderma cadaverica Hym.: Bethylidae 

Scleroderma cadaverica is listed by Herting and Simmonds (1973) as a 

natural enemy of H. hampei, but only some of the specimens before Benoit 

(1957) when he prepared the taxonomic description had been reared from 

that species, others being stated to come from small beetles boring in cane 

furniture. The North American species of Scleroderma are stated by 

Krombein et al. (1979) to be parasitic on the larvae of small wood-boring 

beetles, the female wasps frequently stinging people inhabiting infested 

houses. The African specimens of S. cadaverica were submitted to European 

specialists for identification and description because stinging by females 

(which may be either winged or apterous) had caused severe dermatitis to 

African and European people. No responsible person would consider using 

this insect in biological control projects. 

Comments 

Hypothenemus hampei has established itself in most of the coffee growing 

areas of the world, but there are still uninfested countries, such as Australia, 

Hawaii, Papua New Guinea, Vanuatu and the Solomon Islands. Quarantine 

is of critical importance to these countries and it is important to ensure that 

coffee imported into clean areas has been completely disinfested. Thorough 

drying of the seed coffee is an indispensable supplement to disinfestation 

techniques. 

Although great advances have been made in recent years in the chemical 

control of H. hampei, it would be a great advantage to have the support of 

additional measures. Rhodes and Mansingh (1981) found chemical control 

inadequate on its own in Jamaica and advocated its integration with cultural 

practices and Bardner (1978) emphasised the need to harmonise cultural, 

biological and chemical control of coffee pests. Hernandez Paz and Penagos 

Dardon (1974) found, in Guatemala, that low-volume sprays of endosulfan 

could completely destroy H. hampei in berries on the bushes and, according 

to Mansingh and Rhodes (1983), this chemical is in extensive use in Central 

and South America. However, a very high level of resistance to endosulfan 

in H. hampei is reported in New Caledonia (Brun et al. 1989, 1990), raising 

concern that this valuable insecticide may not remain an effective material 

for long.


Plantation sanitation is an old-established tradition in pest control, and 

the coffee berry borer has long been attacked from this angle. The life cycle 

of the borer, indeed, lends itself to this approach, as it is narrowly specific to 

the coffee berries. In Java, Roepke (1912) and Leefmans (1924b) 

recommended the total destruction of infested or susceptible berries over a 

period long enough to break the life cycle of the coffee berry borer. A period 

of three months was aimed at, although some records of the longevity of 

beetles exceed this. Measures taken involve the collection of all fallen 

berries and the picking of any that may have escaped the harvest, plus the 

continuous removal of all young berries on which adult female beetles might 

feed. Friederichs (1922) and Rutgers (1922) reported successful application 

of the method in Java. The latter reported that, on estates which had applied 

the measures, the percentage of infested berries fell from 40% to 90% to 

between 0.5% and 3.0%. In New Caledonia application of the method 

reduced infestation from 80% to 10% (Cohic 1958). In Malaysia, Corbett 

(1933) recommended picking at weekly intervals (or the shortest period 

practicable, according to size of holding) of all bored green, ripe and 

blackened berries on the bush and from the ground. A host-free period of six 

months was recommended for the eradication of the beetle from isolated 

plantations. In Mexico, Baker (1984) concluded that berries are not infested 

while lying on the ground surface. Nevertheless any infested berries allowed 

to lie where they fell ultimately generated large numbers of beetles. 

Turning to biological control, it is generally possible to gain some idea of 

the likely effectiveness of natural enemies in a new country from their 

impact in their country of origin. In a new country the enemies may be more 

effective if freed from hyperparasites before transfer or less effective if they 

are poorly adapted to their new environment. Based on their evaluation of 

the population dynamics of H. hampei and its parasitoids, Moore and Prior 

(1988) and Murphy and Moore (1990) were optimistic about the value of 

biological control as a key component of successful integrated management 

of H. hampei. Although this indeed seems probable, the reports from Africa 

are far from uniform and the coffee berry borer is a problem in some areas. 

This may be due to whether arabica or robusta coffee is involved and to the 

widely different conditions under which coffee is grown and harvested 

since, for example, parasitoids may be less effective when coffee is shaded 

(Hargreaves 1935). 

Hargreaves (1926, 1935) considered that both Prorops nasuta and 

Heterospilus coffeicola were important in regulating populations of 

H. hampei in commercial coffee at about 1200m in Uganda and Pascalet 

(1939) arrived at a similar view concerning these parasitoids in Cameroon. 

On the other hand, Abasa (1975) in Kenya, De Toledo Piza and Fonseca


(1935) and Ingram (quoted by Le Pelley 1968, p 125) in Uganda and Sladden 

(1934) and Schmitz and Crisinel (1957) in Zaire concluded that parasitoids 

had little influence on the number of bored berries. Murphy and Moore 

(1990) considered that P. nasuta did not have a major impact in Western 

Kenya where H. coffeicola was not encountered. There, coffee berries 

infested with H. hampei, ranged in being attacked by P. nasuta from 0% in 

the wet season to 25% in the dry season and P. nasuta populations did not 

build up until after the annual coffee harvest when populations of H. hampei 

had already crashed from their annual peak. 

Although P. nasuta is present in Ivory Coast (Diro, Man) it is usually 

rare and H. coffeicola apparently absent. However, Cephalonomia 

stephanoderis was common and up to 50% of colonies of H. hampei in black 

berries were parasitised, resulting in important population reduction 

(Ticheler 1961), a conclusion contested by Koch (1973) on unsubstantiated 

grounds. It is relevant that, in the presence of C. stephanoderis, but without 

any chemical treatment, Cochereau and Potiaroa (1994) reported that only 

2% of the coffee beans (4% of the berries) were attacked in Ivory Coast by 

H. hampei. They contrast this with the situation in New Caledonia where it 

was common to find 50% of the berries bored, even in plantations treated 

with endosulfan. 

Experience in biological control suggests that the negative assessments 

of the value of parasites in Africa may reflect the restraining influence of 

hyperparasites (such as Aphanogmus (= Ceraphron) dictynna), predators, 

competitors and diseases, but it seems that no such impediments were taken 

to Brazil with the original stocks of Prorops nasuta. Nevertheless, Le Pelley 

(1968) stated that there was no evidence that, 35 years after the introduction, 

Brazilian coffee growers had to invest less effort in other control measures 

and Yokayama et al. (1978), 45 years after its introduction, gave a 

depressing picture of its impact. Greater success may attend the more recent 

employment of Cephalonomia stephanoderis. 

In spite of the conflicting reports, there are, on balance, good reasons for 

maintaining optimism that natural enemies can play an important role in 

reducing the losses caused by H. hampei. There are the still to be evaluated 

prospects that C. stephanoderis will prove useful and there are various 

strains (and intraspecific crosses) of P. nasuta, in addition to Phymasticus 

coffea, Heterospilus coffeicola and Goniozus to consider. It is quite possible 

also that further work, especially in countries such as Ethiopia, which have 

not been studied, will reveal additional parasitoid species. It is possible that 

nematodes may prove effective in controlling infestations in fallen berries 

and likely that applications of a virulent strain of Beauveria bassiana may 

prove to be a valuable alternative to pesticides. It is fortunate for Southeast


Asian countries that active work on both parasitoids and B. bassiana is in 

progress in a number of South American countries and in New Caledonia, 

from which valuable new information will emerge. However, this is not a 

justification to delay action if H. hampei is high enough on the priority list, 

since it is likely that parasitoid cultures and relevant expertise will be more 

readily and economically available now than they will be some years hence.

4.11 Leucinodes orbonalis 

India 

Myanmar 

++ 

20° 

Laos 

+ 

0° 

20° 

China 

++ 

Thailand 

+ 

Cambodia 

++ 

Vietnam 

+++ 

+++ 

++ Brunei 

Malaysia 

+ 

Singapore 

P 

Indonesia 

Taiwan 

++ 

P 

Philippines 

Australia 

Papua 

New Guinea 

185 

It appears that Leucinodes orbonalis has been introduced from India to Southeast Asia. 

In India and most other countries, egg plant (brinjal) is protected against insect pests with 

heavy applications of broad spectrum pesticides, which must also suppress parasitoids 

and predators. Several parasitoids, and especially the ichneumonid Trathala flavoorbitalis, 

are capable of producing in excess of 50% combined mortality. If levels of this magnitude 

are combined with the widespread adoption of partially resistant egg plant cultivars, there 

are good reasons for believing that substantial pest control would be achieved. 

20° 

0° 

20°


Leucinodes orbonalis GuenŽe 

Rating 

Origin 

Distribution 

Lepidoptera: Pyralidae 

brinjal fruit borer, eggplant fruit and shoot borer 


+++ Viet, Brun, Phil 

18 ++ Myan, Camb, Msia ++ absent 

+ Thai, Laos, Sing 

P Indo 

According to Purseglove (1968), eggplant ( Solanum melongena) 

is native to 

India and there are certainly many varieties in cultivation there. It is 

probably safe to conclude that L. orbonalis is also native to India, although it 

is not confined to eggplant and also occurs widely in Africa (see below). 

Leucinodes orbonalis is closely related to Central American Neoleucinodes 

species such as N. elegantalis, 

which is a widespread pest there of eggplant, 

and other Solanaceae. This suggests that parasitoids of Neoleucinodes may 

be of interest for Leucinodes and vice versa. 

L. orbonalis occurs in the Indian subcontinent (Andaman Is, India, 

Pakistan, Nepal, Bangladesh, Sri Lanka), Southern Asia all 10 Southeast 

Asian nations, also Hong Kong, China, Taiwan, Japan, Africa (Burundi, 

Cameroon, Congo, Ethiopia, Ghana, Kenya, Lesotho, Malawi, 

Mozambique, Nigeria, Rwanda, Sierra Leone, Somalia, South Africa, 

Tanzania, Uganda, Zimbabwe) (CIE 1976; Tamaki and Miyara 1982; 

Whittle and Ferguson 1987; Veenakumari et al. 1995) and is also reported 

from Congo (Dhankar 1988) and Saudi Arabia (FAO 1982). It is not 

recorded from Australasia, Oceania, the Americas or Europe and was 

apparently absent until recently from the Philippines (CIE 1976), although 

Navasero (1983) has now reported it there. Between 1977 and 1987 there 

were 1291 interceptions of L. orbonalis at U.S. ports of entry, most on eggplant 

fruit in passenger baggage (Whittle and Ferguson 1987), and there 

must be significant risks also of its entry to Australia and the Pacific.

Biology 

Host plants 

4.11 


187 

The flat, oval eggs are mostly laid at night, either singly or in groups of 2 to 4 

(and up to 200 per female), on the lower surface of young shoots, flower 

buds and calyces of developing fruits. They hatch in 4 days at an optimum 

temperature of 30¡C and relative humidity of 70% to 90%. Larval 

development occupies 14 days and pupal development 19 days. With a 

preoviposition period of 2 days, this results in a generation time of about one 

month. Details from three authors are shown in Table 4.11.1. Up to 9 larvae 

have been found in a single fruit and, when mature, pupate within a tough 

silken cocoon on the fruit, stem or among ground litter (Tamaki and Miyara 

1982; Khoo et al. 1991; Yin 1993). In the absence of fruit, larvae feed on the 

growing points of the plant. In the plains of India it occurs throughout the 

year but, at higher altitudes, cold weather interrupts its development and it 

overwinters as a larva in a silken cocoon, usually 1 to 3 cm below the soil 

surface. It is capable of surviving temperatures as low as Ð6.5¡C (Lal 1975). 

It thrives best under warm, moist monsoonal conditions. L. orbonalis can be 

reared in the laboratory on dried eggplant fruit or on a semi-synthetic diet 

(Islam et al. 1978, Patil 1990). Virgin females produce a pheromone that 

attracts males (Gunawardena 1992; Yasuda and Kawasaki 1994). 

In addition to eggplant, which is its main host, L. orbonalis is reported to 

feed on several other Solanum species, e.g. S. tuberosum (potato: shoots 

only) (Fletcher 1916; Mehto et al. 1980; Isahaque and Chaudhuri 1983), 

S. nigrum (black berry nightshade) (Nair 1967; Das and Patnaik 1970; 

Isahaque and Chaudhuri 1983 ), S. indicum, 

S. myriacanthum (shoots only) 

(Menon 1962; Isahaque and Chaudhuri 1983) and S. xanthocarpum (Menon 

1962). It has also been reported from tomato ( Lycopersicon esculentum) 

(Hargreaves 1937; Das and Patnaik 1970), potato ( Solanum tuberosum) 

and 

several unexpected plants, including cape gooseberry (Pillai 1922), green 

pea pods (Hussain 1925), mango shoots (Hutson 1930), cucumber, sweet 

potato and capsicum (Whittle and Ferguson 1987). Screening egg plant 

varieties for resistance to L. orbonalis has revealed several that are relatively 

resistant. Thick-skinned varieties appear to be more resistant (Patil and Ajri 

1993; Patel et al. 1995).

Table 4.11.1 

Development times, and other life history data (rounded) of Leucinodes orbonalis 

Reference Atwal and Verma 1972 Baang and Corey 1991 Mehto et al. 1983 

Temperature 20¡C 25¡C 30¡C 35¡C 

egg development(days) 9 6 4 3 5 6 

egg survival (%) 63 69 78 55 

larval development(days) 29 20 14 12 18 15 

larval survival (%) 53 72 69 48 

pupal development(days) 17 13 9 7 10 12 

pupal survival (%) 57 65 71 55 

longevity females(days) 11 7 6 3 3 8 

males(days) 9 7 4 2 2 4 

eggs/female 188 225 248 86 85Ð254 122 


Damage 

4.11 


189 

All stages of eggplant are attacked by L. orbonalis, 

which is regarded as one 

of its major insect pests. Larvae bore into the tender shoots of both seedlings 

and after transplantation older plants, causing wilting and death of the 

growing tips. Later, they bore into flower buds and fruits. The damaged buds 

are shed and the fruits carry circular holes, sometimes plugged with frass. 

Such fruits are unmarketable. The yield loss varies with location and season 

and is greatest when temperature and humidity is high. Losses range from 20 

to 60% (Krishnaiah 1980; Dhankar 1988, Roy and Pande 1994) or even 

higher (Akhtar and Khawaja 1973; Lal 1991). It is reported that Vitamin C in 

bored fruit can be reduced by 60% (Hami 1955). 

There is an extensive literature dealing with the screening, mainly in 

India, for resistance of eggplant cultivars to L. orbonalis. 

Some cultivars are 

far less damaged than others, although no information is available on the 

genes involved. The less susceptible cultivars generally have one or more 

morphological characteristics, including compact vascular bundles in a thick 

layer, lignified cells and less area of pith in the shoots, tougher fruit skin and 

a tight calyx to hinder larval entry. Biochemical factors involved include a 

low protein and sugar content in resistant genotypes and a higher silica and 

crude fibre content in the shoots, which adversely affects growth rate, pupal 

period, survival, sex ratio and fecundity. The wild relatives of Solanum 

melongena that are not attacked by L. orbonalis often have a high alkaloid 

content, which may be responsible, but this attribute would not be desirable 

in an edible product (Dhankar 1988). 


Table 4.11.2 lists the natural enemies of L. orbonalis. 

It is striking that most 

are from India and Sri Lanka, that the records from Malaysia and the 

Philippines are the only ones from Southeast Asia and that the species there 

have not been reported elsewhere. Although it is possible that some of the 

Indian and Sri Lankan species have a restricted distribution, the lack of 

records from elsewhere probably means that the necessary investigations 

have not been carried out.

Table 4.11.2 

Natural enemies of Leucinodes orbonalis 

Species Stage attacked % parasitisation Country Reference 

NEUROPTERA 

CHRYSOPIDAE 

Chrysopa kulingensis 

DERMAPTERA 

egg 

larva 

12.5 

2-4 

China Yang 1982 

UNIDENTIFIED (15) Philippines Navasero 1983 

DIPTERA 

SARCOPHAGIDAE 

Amobia sp. 

TACHINIDAE 

Malaysia Thompson 1946 

Pachyophthalmus sp. Malaysia Corbett 1929; Yunus & Ho 1980 

Pseudoperichaeta sp. 5.7 India Patel et al. 1971 

Sturmia parachrysops 

India Thompson 1946 

UNIDENTIFIED 

HYMENOPTERA 

BRACONIDAE 

pupa 5Ð8 China Yang 1982 

Apanteles sp. 3.1-11.1 Philippines Navasero 1983 

Bracon greeni 

larva; ecto lab only India Venkatraman et al. 1948 

Bracon sp. larva; ecto 9.2Ð28.1 India Tewari & Sardana 1987a 

Campyloneura sp. larva 1Ð2 India Tewari & Moorthy 1984 

Chelonus sp. 

15.5 

Phanerotoma sp. 4.6 

7.4 

Phanerotoma sp. nr. 

hindecasisella 

Philippines 

Sri Lanka 

India 

Sri Lanka 

Navasero 1983 

Sandanayake & Edirisinghe 1992 

Patel et al. 1971 


larva 1Ð2 India Patel et al. 1971; 

Tewari & Moorthy 1984 


Table 4.11.2 (contÕd) Natural enemies of Leucinodes orbonalis 

Species 

HYMENOPTERA 

Stage attacked % parasitisation Country Reference 

CHALCIDIDAE 

Brachymeria lasus 

Philippines Navasero 1983 

Brachymeria sp. 

EULOPHIDAE 


Dermatopelte 

(= Dermatopolle) 

sp. 

ICHNEUMONIDAE 

pupa 16 China Yang 1982 

Cremastus hapaliae 


Eriborus argenteopilosus larva 0.5Ð2 India Tewari & Sardana 1987b 

Itamoplex sp. pupa 9Ð15 India Verma & Lal 1985 

Pristomerus testaceus 

larva; ecto India Ayyar 1927 

Trathala flavoorbitalis 

Trathala striata 

Xanthopimpla punctata 

BACTERIUM 

BACULOVIRUS 

larva 36.2 

3.6Ð9.1 

12.9Ð18.2 

Sri Lanka 

India 

India 

India 

Malaysia 


Mallik et al. 1989 

Naresh et al. 1986a, b, 

Patel et al. 1967 

Yunus & Ho 1980 

Malaysia C.L. Tan pers. comm. 1994 


larva 2Ð3 China Yang 1982 

larva 1.1Ð6.4 India Tewari & Singh 1987 

4.11 


191


CHINA 

The life history of L. orbonalis was studied by Yin (1993), in Hunan where 

up to 6 generations were completed annually and overwintering occurred in 

the pupal stage. Yang (1982) recorded two pupal parasitoids, the 

polyembryonic wasp Dermatopelte sp., causing 16% parasitisation (and 

producing 12 to 21 individuals per host pupa) and an unidentified dipteran 

causing 5% to 8% mortality. A predator, Chrysopa kulingensis consumed 

12.5% of eggs and 2% to 4% of larvae. A disease, presumably of bacterial 

origin, killed 2% to 3% of larvae. 

INDIA, SRI LANKA 

The ichneumonid Trathala flavoorbitalis appears to be the most effective 

parasitoid so far recorded, with an average parasitisation rate in Sri Lanka of 

36.2% (Sandanayake and Edirisinghe 1992). In Hanyana, India, Naresh et al. 

(1986a) recorded rates from 12.9% to 18.2% and in Bihar, Mallik et al. 

(1989) 3.6% to 9.1%. These were higher than the combined rates of 1% to 

2% by Phanerotoma sp. and Campyloneura sp.. T. flavoorbitalis is a very 

widespread species and attacks the larvae of many species of Lepidoptera. 

Bracon sp. from near Bangalore, India, with a parasitisation rate ranging up 

to 28.1% (Tewari and Sardana 1987a), Chelonus sp. ranging up to 5.5% 

(Sandanayake and Edirisinghe 1992) and Itamoplex sp. up to 15% (Verma 

and Lal 1985) are all capable of producing significant mortality. 

PHILIPPINES 

A dermapteran larval predator and 5 parasitoids (the braconids Apanteles sp. 

(on larvae) and Chelonus sp. (on pupae), the chalcidids Brachymeria lasus 

(= B. obscurata) and Brachymeria sp. (larvae and pupae), and the 

ichneumonid Xanthopimpla punctata) were found attacking L. orbonalis in 

the field. Apanteles sp. and the dermapteran were the most abundant 

(Navasero 1983). It appears that L. orbonalis has only been recognised in the 

Philippines since the early 1970s. 


There have been none. 


Bracon sp. Hym.: Braconidae 

This larval ectoparasitoid was found near Bangalore, India attached to the 

thorax of the host larva. It pupated in a silk cocoon inside the tunnel made by 

its host and caused parasitisation ranging from 9.2% to 28.1%. It was 

regarded as a promising parasitoid (Tewari and Sardana 1987a).

4.11 Leucinodes orbonalis 193 

Itamoplex sp. Hym.: Ichneumonidae 

Adult Itamoplex sp., 8 to 10 mm in length, were reported from Kulu Valley, 

Himachal Pradesh, India where the winter temperature drops as low as Ð8¡C. 

L. orbonalis overwinters in the larval (?prepupal) stage in an earthen cocoon 

attached to the host plant, usually 1 to 3 cm below the soil surface. The 

parasitoid emerged from 9% to 15% of these cocoons. Itamoplex (= Cryptus) 

sp. is recorded attacking a range of host Lepidoptera in cocoons (Verma and 

Lal 1985). 

Trathala (= Cremastus) flavoorbitalis Hym.: Ichneumonidae 

This is a widespread, non-specific parasitoid of lepidopterous larvae. It 

occurs naturally in India, Japan, Myanmar, Sri Lanka, the Philippines and 

Singapore and has been established in Canada, Hawaii and continental USA 

for biological control of several important lepidopterous pests. It is recorded 

from at least 5 families of Lepidoptera, involving over 40 different hosts, 

most of them pest species (Bradley and Burgess 1934). It is not known 

whether there are strains that prefer to attack particular hosts. 

T. flavoorbitalis is recorded from L. orbonalis in India and also in Sri 

Lanka where L. orbonalis is its major host and where an average 

parasitisation level of 36.2% is reported (Sandanayake and Edirisinghe 

1992, 1993). In Hissar, India, Trathala was the only parasitoid of 

L. orbonalis, with levels of attack on larvae ranging from 13.2% to 18.2% in 

winter to 12.9% in summer at a time when 95.2% of fruit was infested 

(Naresh et al. 1986a, b). 

T. flavoorbitalis females commence ovipositing 4 days after emergence, 

with a preference for 3rd, 4th and 5th instar host larvae. In the laboratory, 

only a fraction of 1st instar larvae are stung and all soon die from the 

encounter, a fate shared by about half the 2nd instar larvae that are stung. In 

later instars 68Ð91% were stung, but without early mortality. Not all of these 

received eggs, although some received up to 5, with only one parasitoid larva 

developing beyond the first instar. Parasitoid development time from egg to 

adult was about 23 days at 28¡C. Successful development occurred in 53% 

of the 3rd, 57% of the 4th and 41% of 5th instar host larvae, adult wasps 

emerging after pupation of the host. In the field, the parasitoid attacks the 

host larva by inserting its ovipositor into the hole bored into the fruit and 

through which the larva pushes out frass (Bradley and Burgess 1934; Naresh 

et al. 1986a; Sandanayake and Edirisinghe 1992, 1993).


Comments 

It is not clear how many of the natural enemies are sufficiently host specific 

to be confidently transferred as biological control agents although, where 

alternative hosts are known, these are also pests. Thus, Eriborus 

argenteopilosus has a wide host range, including Condica (= Prospalta) 

capensis, which attacks safflower and sunflower, Helicoverpa armigera, 

and Spodoptera exigua (Tewari and Sardana 1987a,b). The ichneumonid 

Pristomerus testaceus has been bred from the brinjal stem-borer Euzophera 

ferticella (Ayyar 1927). It is not known whether the ichneumonid 

Phanerotoma sp. nr hindecasisella is a distinct species. True 

P. hindecasisella has been reported from several lepidopterous families in 

India or Sri Lanka: Gelechiidae (Dichomeris eridantis), Noctuidae (Earias 

insulana), Pyralidae (Eutectona (= Pyrausta) macheralis, Hendecasis 

duplifascialis, Maruca vitrata (= M. testulalis), Nephopterix rhodobasalis, 

Syllepte derogata) and Tortricidae (Leguminivora (= Cydia) ptychora) 

(Thompson 1953; Fellowes and Amarasena 1977; Kumar et al. 1980; 

Tewari and Moorthy 1984). 

Bracon greeni is best known as a primary ectoparasitoid of the 

lepidopterous lac predator Eublemma amabilis (Noctuidae) and has not been 

reported to parasitise any other host in nature. However, under laboratory 

conditions, it was successfully reared on L. orbonalis (Venkatraman et al. 

1948). 

Although entomopathogenic nematodes have not been recorded 

attacking L. orbonalis in the field, Steinernema (=Neoaplectana) 

carpocapsae (DD136 strain) produced 73.3% mortality of larvae in the 

laboratory in 72 hours (Singh and Bardhan 1974). 

The weight of evidence suggests that L. orbonalis originated in India and 

spread into Southeast Asia. It is most surprising that it has only 

comparatively recently become a pest Ñ and a serious one Ñ in the 

Philippines which, in 1990, had 16 000ha under egg plant and produced 

113 000 tonnes, second only to Indonesia in production in Southeast Asia 

(FAO 1991). In view of the steady spread around the world of so many other 

pests it is also surprising that Australia, the Pacific, the Americas and Europe 

are still free from L. orbonalis. 

It might well be assumed that not all of its natural enemies in India have 

accompanied it during its spread. However, reports of high damage levels to 

susceptible egg plant cultivars in India do not provide much confidence that 

the natural enemies there are particularly effective, unless their efficacy is, 

perhaps, reduced by insecticide applications or so-far-unreported 

hyperparasitoids.

4.11 Leucinodes orbonalis 195 

Nevertheless, parasitisation in Sri Lanka is quite impressive, with 

Trathala flavoorbitalis averaging 36.2%, Chelonus sp. 15.5% and 

Phanerotoma sp. 7.4%. The combined average rate of 59.1% would 

certainly be capable of causing an important lowering of moth populations, 

particularly if associated with the high levels of host plant resistance that are 

available. It is probable that the widespread high rates of application of 

broad spectrum insecticides currently used are preventing natural enemies 

from exerting much effect. An investigation of what natural enemies of 

L. orbonalis are already present in Southeast Asia is required to enable a 

decision on the attractiveness of this pest as a biological control target. It is 

probably safe to conclude that T. flavoorbitalis, the most effective parasitoid 

so far reported, is present in most, if not all, of Southeast Asia, but it would 

be relevant, in relation to possible host-preferring strains, to determine 

whether it attacks L. orbonalis throughout the region. 

It is interesting that the closely related South American Neoleucinodes 

elegantalis has a quite different suite of parasitoids. Of 2500 larvae collected 

in the field 1.6% were parasitised by the encyrtid Copidosoma sp. and 0.08% 

by the tachinid Lixophaga sp. Of 527 pupae, 0.38% were parasitised by the 

ichneumon Calliephialtes sp. The fungus Beauveria sp. caused 55% 

mortality. Trichogramma sp. parasitised 89% of eggs laid on egg plant and 

tomato (Plaza et al. 1992). Of these species, Beauveria and the 

Trichogramma sp. might be relevant to the Southeast Asian scene.

4.12 Nezara viridula 

India 

Myanmar 

++ 

20° 

Laos 

+ 

0° 

20° 

China 

++ 

Thailand 

+ 

Cambodia 

+ 

Vietnam 

++ 

P 

+ Brunei 

Malaysia 

+ 

Singapore 

+ 

Indonesia 

Taiwan 

++ 

P 

Philippines 

Australia 

Papua 

New Guinea 

++ 

197 

Nezara viridula is probably native to the Ethiopian region, but is now dispersed widely 

throughout the warmer regions of the world. 

There have been a number of major successes with biological control of the green 

vegetable bug, particularly with an egg parasitoid, Trissolcus basalis. 

Control has been 

supplemented in Hawaii by two parasitoids of adults, Trichopoda pilipes and T. pennipes, 

which, however, have failed to establish in most other places. 

The main areas where biological control using T. basalis has been unsuccessful are 

often associated with extensive plantings of soybean. It has been suggested that the 

surface hairyness of most soybean cultivars reduces the effectiveness of this parasitoid. If 

this proves to be correct the way is open for the selection of cultivars with this attribute 

modified. 

Over 80 parasitoids of N. viridula are known, amongst which there are several 

potentially valuable, untried species that are clearly worthy of investigation. 

N. viridula is an attractive target for biological control, especially where it is a problem in 

an area not closely associated with extensive soybean plantings. 

20° 

0° 

20°


Nezara viridula (Linnaeus) 

Rating 

Origin 

Hemiptera, Pentatomidae 

green vegetable bug, southern green stink bug (USA) 


10 ++ Myan, Viet ++ 14 ++ Cook Is, Fr P, Niue, 

PNG, Sam 

+ Thai, Laos, Camb, 

Msia, 

Distribution 

+ Fiji, Kiri, N Cal, 

A Sam 

Sing, Indo 

P Brun, Phil P FSM, Guam, Sol Is, 

Tong, Van 

This account updates the chapter on Nezara viridula in Waterhouse and 

Norris (1987), and the valuable account of Jones (1988), in relation to 

prospects for biological control. 

The locality of the holotype Ôin IndiisÕ (Linnaeus 1758) has been interpreted 

as India or the East Indies. However it may well have been brought there by 

European travel since an analysis of genetic colour variants and the relative 

abundance of fairly specific insect parasitoids led Hokkanen (1986) and 

Jones (1988) to the conclusion that the original home of N. viridula was in 

the Ethiopian (Afrotropical) or Mediterranean region, rather than in 

Southeast Asia (Yukawa and Kiritani 1965). Furthermore, Africa is 

considered to be the centre of the genus Nezara (Freeman 1940). 

Azores, Canary Is, Bermuda, the Mediterranean littorale, most of Africa and 

the Middle East, Madagascar, Mauritius, Reunion, Rodriguez, Seychelles, 

Asia (exclusive of desert areas and those with very cold winters) Korea and 

southern Japan, Southeast Asia, Papua New Guinea (including Bismarck 

Archipelago), most of the oceanic Pacific nations (Butcher 1981), Australia, 

New Zealand, Hawaii, southern USA, Mexico and other Central American 

countries, West Indies generally, and much of South America (Anon. 1970). 

It is not known to occur in Tokelau, Tuvalu or the Marquesas (Waterhouse 

1985, 1997).

Biology 

Damage 

4.12 


199 

The entirely green colour form smaragdula of N. viridula is the one that 

occurs widely in the Pacific area. In other areas of the world it is 

accompanied by several other colour forms (Yukawa and Kiritani 1965). 

The barrel-shaped eggs are usually laid at night in neat rafts, commonly of 

80 to 120 or more eggs and they are cemented firmly to one another and to 

the sheltered surface of a leaf. The eggs hatch in 4 to 9 days, and the newly 

emerged nymphs remain together near the eggshells for a day or two, a 

degree of gregariousness persisting also in the next instar or two. There are 

five nymphal stages before the adult emerges, 24 to 60 days after hatching, 

depending on temperature. At 25Ð28¡C, 55Ð65% RH and 14 hours daylight 

the development periods (in days) were: egg, 4.8; 1st instar nymph, 3.8; 2nd 

instar, 5.2; 3rd instar, 4.5; 4th instar, 6.4; 5th instar, 11.9 (Harris and Todd 

1980). The nymphal stages are multicoloured, but the adults are a uniform 

green in the form smaragdula. 

The adults can fly strongly. There may be 

four generations in a year in coastal New South Wales, and perhaps more in 

areas with no perceptible winter. 

The adult bugs live up to 3 weeks in hot weather. In regions with a cold 

winter those of the autumn generation may live much longer, hibernating in 

debris, under bark, or in buildings, inactive and non-reproductive. Such 

hibernating bugs change colour from green to brown. In areas with a less 

severe winter the still-green adults may remain active, although nonreproductive. 

Waite (1980) showed that the adults and nymphs tend to bask 

exposed on the surface of the plant canopy in the early daylight 

hoursÑbehaviour that can be availed of in applying chemical control 

measures (Kamal 1937; Clausen 1978; Hely et al. 1982; Singh and Rawat 

1982; Todd 1989). 

When not controlled by chemicals or natural enemies, this bug can be a 

serious pest of a very wide range of crops and ornamental plants. In Australia 

there are recommendations for its chemical control on beans, cucurbits, 

peas, potatoes, tomatoes, passionfruit, groundnuts, sorghum, soybeans, 

sunflowers and tobacco (Anon. 1967, 1979b; Miller et al. 1977), but it also 

attacks maize, crucifers, spinach, lucerne and many other legumes, grapes, 

oranges and many other fruits and seeds, and macadamia (Ironside 1979; La 

Croix 1986) and pecan nuts (Seymour and Sands 1992). Undoubtedly there 

would be recommendations for a much wider range of cultivated plants were 

it not for the fact that biological control is effective now in many situations


(Anon. 1967; Hely et al. 1982). In other countries it can be a pest of rice, 

sesame and other grains, lablab, guavas, cowpeas, capsicums, cotton and 

many other plants. It also infests and breeds on many weeds, which afford it 

harborage. 

The attack of the bugs is concentrated chiefly on fruits and fruiting 

bodies, which, through the removal of sap and the injection of saliva, show 

discoloration, malformation, stunting and shrivelling. Heavily attacked 

tomatoes, for instance, are repulsive and inedible and even lightly attacked 

ones are unmarketable. Passlow and Waite (1971), Goodyer (1972) and 

Romano and Kerr (1977) attribute serious losses in Australian soybean crops 

to the attack of this pest. Miller et al. (1977) measured the yield reduction on 

soybeans and showed that an important effect of the bugs feeding was a 

reduction in the germinability of seed. On soybean in Brazil, Corso et al. 

(1975) showed that green vegetable bug attack affected pod development, 

increased pod fall, and reduced the number of seeds per pod. Link et al. 

(1973) demonstrated reductions in germination percentage and oil content 

and an increase in relative protein content in heavily damaged seeds. A list of 

references dealing with N. viridula and its association with soybeans has 

been prepared by DeWitt and Godfrey (1972). 

Some cultivars of soybeans are less damaged by N. viridula than others 

and a strain of soybean that appears to have a high level of resistance has 

been selected (Gilman et al. 1982). Mild antibiosis and non-preference were 

factors contributing to resistance (Kester et al. 1984). The adverse effects of 

the genotype (PI 171444) on the biology of a parasite Telenomus chloropus 

attacking the eggs of N. viridula feeding on it have been studied by Orr et al. 

(1985b). 

The green vegetable bug has been shown to carry spores of fungal 

diseases from plant to plant (Corso et al. 1975) and to transmit plant 

pathogens during feeding (Kaiser and Vakili 1978). 


Introduced parasitoids have brought about successful biological control of 

this pest in a number of countries and there is an extensive literature on the 

subject. Nine species of parasite attacking Nezara viridula were listed by 

Thompson (1944), 27 by Hokkanen (1986) and 57 by Jones (1988). At least 

eighty are now included in Table 4.12.1 which is a modification and 

extension of that in Jones (1988). 

The species belong to two families of Diptera and six families of 

Hymenoptera. It is notable from the entries that relatively little is known of 

the parasitoids of N. viridula in the Ethiopian or Mediterranean regions,

4.12 


201 

which constitute the presumed area of origin of the green vegetable bug. Egg 

parasitoids are the most numerous and all are Hymenoptera, whereas 

nymphal and adult parasitoids are, with one exception, all Diptera. Two 

hyperparasitoids of the major egg parasitoid Trissolcus basalis are known 

from Australia, (both are species of Acroclissoides: 

Clarke and Seymour 

1992) and one hyperparasitoid of the fly Trichopoda pennipes from Hawaii 

( Exoristobia philippinensis: 

Davis and Krauss 1965). Predators are not dealt 

with as all are known to be, or suspected as being, widely polyphagous and 

hence unlikely to be approved for introduction to new areas. Nevertheless, 

they play a significant role in maintaining N. viridula populations at low 

levels. For example, in one study in soybeans in Louisiana, Stam et al. 

(1987) found 18 insect and 6 spider species to be predators and that they 

were responsible for 33.6% of the total mortality of N. viridula that occurred 

from egg to adult. 

The Scelionidae is the most important of the six families of 

hymenopterous egg parasitoids. In it Trissolcus basalis is not only the most 

important species, but also the most widespread. It attacks the eggs of a 

number of other pentatomids, but appears to have a preference for 

N. viridula: 

it is the dominant parasitoid of N. viridula eggs wherever it 

occurs. Many other species are listed, in particular in the genera Trissolcus, 

Telenomus, Ooencyrtus and Gryon, 

but a number appear to have no close 

relationship with N. viridula and are unlikely to be of value as potential 

biological control agents. Some drought-resistant species from the 

Mediterranean may prove useful. The better known of the more promising 

species are discussed later. 

The tachinid parasitoids of adult N. viridula also oviposit on the cuticle 

of 4th and 5th instar nymphs. Often these eggs are shed before hatching 

along with the cuticle at moulting but, if hatching and penetration of the bug 

occurs, the parasitoid larva matures in the adult. More of the species of 

tachinids are native to South America than elsewhere. They were clearly 

dependent upon other pentatomids before the arrival of N. viridula, 

but 

several now appear to have a preference for it. The most promising of the 

three tachinid parasitoids occuring in the Ethiopian region is the widespread 

Bogosia antinorii, 

which is known only from N. viridula (van Emden 1945; 

Barraclough 1985). 

A picorna-like and a toti-like virus are known from N. viridula 

(Williamson and Wechmar 1992, 1995).

Table 4.12.1 

Parasitoids of the green vegetable bug, Nezara viridula (modified after Jones 1988) 

Parasitoid Geographic range Known host relations Selected references 

DIPTERA 

SARCOPHAGIDAE 

Sarcodexia innota 

Sarcodexia sternodontis 

TACHINIDAE 

Bogosia antinorii 

Cylindromyia rufifemur 

Ectophasia crassipennis 

Ectophasiopsis arcuata 

Euclytia flava 

Gymnosoma clavata 

Gymnosoma kuramanum 

Gymnosoma rotundata 

Trichopoda giacomellii 

(= Trichopoda nigrifrontalis, 

= T. gustavoi 

= Eutrichopodopsis nitens) 

Trichopoda lanipes 

Trichopoda pennipes 

Southern USA Two records ex N. viridula; 

wide host range as primary 

parasitoid and scavenger 

Widespread in Africa Recorded only ex N. viridula 

Australia One record ex N. viridula 

Italy Bred ex N. viridula 

Chile Well adapted to N. viridula 

Drake 1920; Temerak & 

Whitcomb 1984 

Hokkanen 1986 

Greathead 1966, 1971; 

Barraclough 1985 

Cantrell 1984 

Colazza & Bin 1995 

Jones 1988 

USA Generalist parasitoid Aldrich 1995 

Palaearctic, Israel One record ex N. viridula 

Herting 1960 

Japan Takano 1956 

Palaearctic Wide host range; attacks Nezara spp. in Japan Kiritani et al. 1963; Kiritani 

& Sasaba 1969 

Argentina, Brazil, Well adapted to N. viridula 

Blanchard 1966; Mallea et 

Colombia, Paraguay 

al. 1968; Gastal 1977a,b; 

Liljesthršm 1980, 1981; 

Ferreira 1984 

Southern USA One record ex N. viridula; 

attacks other species Drake 1920 

North America, Hawaii Well adapted to N. viridula 

Drake 1920; Todd & Lewis 

1976; Jones 1979; 

Buschman & Whitcomb 

1980; McPherson et al. 

1982 


Table 4.12.1 (contÕd) Parasitoids of the green vegetable bug, Nezara viridula (modified after Jones 1988) 

Parasitoid 

DIPTERA 

Geographic range Known host relations Selected references 


Trichopoda pilipes 

West Indies, Hawaii Well adapted to N. viridula 

Trichopoda sp. Uruguay One record ex N. viridula 

Myers 1931; Nishida 1966; 

Davis 1967 

Guido & Ruffinelli 1956 

HYMENOPTERA 

EULOPHIDAE 

Pleurotropitiella albipes 

EURYTOMIDAE 

Argentina Esquivel 1950 

Neorileya sp. 

EUPELMIDAE 

Brazil Recorded only ex N. viridula Ferreira 1981, 1984, 1986 

Anastatus bifasciatus Italy Bred ex N. viridula Colazza & Bin 1990 

Anastatus dasyni Malaysia Pentatomidae, Coreidae; described ex N. viridula van der Vecht 1933 

Anastatus japonicus East Asia Lepidoptera, Heteroptera; produces only males in 

N. viridula 

Hokyo et al. 1966b; Kiritani 

& Sasaba 1969 

Anastatus sp. Thailand Emerged ex imported eggs of N. viridula Jones 1988 

Anastatus sp. Southern USA Two records ex N. viridula Jones 1988 

Anastatus sp. Australia 

(Queensland) 

One record ex N. viridula Seymour & Sands 1993 

Unidentified sp. 

ENCYRTIDAE 

Hawaii ex N. viridula on macadamia Jones 1992 

Hexacladia hilaris USA One record ex N. viridula Buschman & Whitcomb 

1980 

Ooencyrtus californicus California ex N. viridula and other pentatomids Hoffmann et al. 1991 

4.12 


203


Parasitoid 

HYMENOPTERA 



Ooencyrtus fecundus Morocco VoegelŽ 1961 

Ooencyrtus johnsoni California ex N. viridula and other pentatomids Hoffmann et al. 1991 

Ooencyrtus malayensis Malaysia, Philippines Pentatomidae, Coreidae, Lepidoptera van der Vecht 1933; Jones 

et al. 1983 

Ooencyrtus nezarae East Asia Coreidae, Pentatomidae, Plataspidae; not uncommon 

on N. viridula in Japan 

Hokyo & Kiritani 1966 

Ooencyrtus pityocampae Italy Breeds in eggs of N. viridula and other pentatomids in 

the laboratory 

Tiberi et al. 1991 

Ooencyrtus submetallicus West Indies, Central 

and South America 

Pentatomidae, Coreidae Gahan 1927; Lee 1979; de 

Santis 1985; Ferreira 1986 

Ooencyrtus trinidadensis West Indies, Argentina Pentatomidae, Coreidae Davis & Krauss 1963; 

Davis 1967; de Santis 1985 

Ooencyrtus sp. Brazil One record ex N. viridula Ferreira 1986 

Ooencyrtus sp. Thailand Emerged ex imported eggs of N. viridula Jones 1988 

Ooencyrtus sp. Philippines Possibly is O. malayensis Davis 1967; Corpuz 1969 

Ooencyrtus sp. France One record; recovered ex other pentatomids Jones 1988 

Ooencyrtus sp. Italy Bred ex N. viridula Colazza & Bin 1995 

Ooencyrtus sp. (spp.?) Southern USA Taxonomy and host range not known Drake 1920; Buschman & 

Whitcomb 1980; 

Jones 1988 

Xenoencyrtus hemipterus 

(= X. niger) 

Australia Seymour & Sands 1993 

Xenoencyrtus rubricatus Australia Described ex N. viridula Riek 1962 

Xenoencyrtus sp. Australia Bred ex N. viridula Forrester 1979 



Parasitoid 

HYMENOPTERA 


PTEROMALIDAE 

Pteromalus sp. Egypt One record ex N. viridula Adair 1918 

3 spp. Brazil Ferreira & Moscardi 1995 

SCELIONIDAE 

Gryon fulviventris Africa, Asia Pentatomoidea; attacks N. viridula only in Thailand Anderson 1919; Dry 1921, 

Jones 1988 

Gryon japonicum Japan, Brazil native of Japan Kishino & Teixeira 1994 

Gryon obesum Southern USA, Brazil Records ex N. viridula; attacks other pentatomids Buschman & Whitcomb 

1980; Masner 1983; 

Hoffmann et al. 1991; 

Correa & Moscardi 1995 

Gryon sp. Australia Minor attack on N. viridula Titmarsh 1979 

Gryon sp. Laos One record ex N. viridula; may be G. fulviventris Grist & Lever 1969; Dean 

1978a,b 

Gryon sp. India One record ex N. viridula, may be G. fulviventris Yadava et al. 1982 

Psix lacunatus Asia, Australia Pentatomidae, Scutelleridae; ex N. viridula in Pakistan Johnson & Masner 1985 

Psix striaticeps Africa, India 

Togo 

Pentatomidae; recorded once ex ÔNezaraÕ 

Common on N. viridula and 2 other pentatomids 

Telenomus chloropus Palearctic Pentatomidae; major parasitoid of Nezara spp. in 

E. Asia; females only in Japan 

Fouts 1934; Johnson & 

Masner 1985; 

Poutouli 1995 

Kiritani & Hokyo 1962; 

Hokyo & Kiritani 1963; 

Johnson 1984a 

Telenomus comperei Philippines Cadapan & Alba 1987 

Telenomus cristatus Southern USA, West 

Indies 

Known only ex N. viridula and Acrosternum hilare Johnson 1984a; Orr et al. 

1986 

4.12 Nezara viridula 205


Parasitoid 

HYMENOPTERA 


SCELIONIDAE (contÕd) 

Telenomus cyrus Java, Philippines, 

Taiwan 

Descr. ex N. viridula; host relations unknown Nixon 1936; Taiwan 

Agricultural Research 

Institute 1984; Jones 1988 

Telenomus gifuensis East Asia Pentatomidae, Coreidae; not well adapted to N. viridula Hidaka 1958; Hokyo & 

Kiritani 1963 

Telenomus mormideae South America Attacks N. viridula and other pentatomids Ferreira 1986; Liljestršm & 

Bernstein 1990 

Telenomus pacificus Philippines Cadapan & Alba 1987 

Telenomus podisi North and South 

America 

Pentatomidae; not well adapted to N. viridula Buschman & Whitcomb 

1980; Correa & Moscardi 

1995; Orr et al. 1985a, 

1986 

Telenomus seychellensis East Africa Attacks other spp.; may be common on N. viridula Nixon 1935; 

Croix & Thindwa 1986 

Telenomus sp. Argentina 

Vietnam 

Minor attack on N. viridula eggs Liljestršm & Bernstein 1990 

van Lam 1996 

Trissolcus aloysiisabaudiae East Africa Reportedly common on N. viridula in cotton Fouts 1930; Chiaromonte 

1931; Paoli 1933 

Trissolcus basalis N. & S. America, S. 

Europe, Africa, Hawaii, 

Australia, New 

Zealand, Fiji 

Most important parasitoid of N. viridula outside central 

Africa and eastern Asia 

Miller 1928; Kamal 1937; 

Lever 1941; Buschman & 

Whitcomb 1980; Ferreira 

1980; Orr et al. 1986; 

Colazza & Bin 1995 

Trissolcus brochymenae N. and S. America Recorded ex N. viridula; attacks other pentatomids Johnson 1984b 



Parasitoid 

HYMENOPTERA 


SCELIONIDAE (contÕd) 

Trissolcus crypticus Pakistan ex Plautia crossota and Acrosternum gramineum; bred Clarke 1993a 

readily in N. viridula in lab. in Australia and Hawaii 

Trissolcus euschisti California ex N. viridula and other pentatomids Hoffmann et al. 1991 

Trissolcus hullensis North America, 

Venezuela 

Recorded ex N. viridula; attacks other pentatomids Johnson 1985 

Trissolcus lepelleyi Central Africa Descr. ex N. viridula, an apparently common host Nixon 1936; Le Pelley 1979 

Trissolcus lodosi Turkey Descr. ex N. viridula; nothing else known Szab— 1981 

Trissolcus maro Southern Africa N. viridula is only known host Nixon 1935; Croix & 

Thindwa 1986 

Trissolcus mitsukurii Japan Important parasitoid of N. viridula in Japan Kiritani & Hokyo 1962; 

Hokyo & Kiritani 1963 

Trissolcus oenone Australia Johnson 1991 

Trissolcus ogyges Australia one recent record Seymour & Sands 1993 

Trissolcus rudus Vietnam van Lam 1996 

Trissolcus scuticarinatus South America One record ex N. viridula; attacks other pentatomids Ferreira 1986 

Trissolcus sipius East Africa Descr. ex N. viridula but not reported since Nixon 1936 

Trissolcus solocis Florida, Mexico Recorded ex N. viridula; attacks other pentatomids Buschman & Whitcomb 

1980; Johnson 1985 

Trissolcus thyantae Eastern N. America Recorded ex N. viridula; attacks other pentatomids Johnson 1985 

Trissolcus urichi Brazil Ferreira & Moscardi 1995 

Trissolcus utahensis California ex N. viridula and other pentatomids Hoffmann et al. 1991 

Trissolcus sp. Taiwan 

India 

Recorded ex N. viridula; host relations unknown 

Recorded ex N. viridula 

Taiwan Agricultural 

Research Institute 1984; 

Nath & Dutta 1994 



The role of pheromones and other chemical 

secretions 

Sexually mature male N. viridula release a pheromone that is a powerful 

attractant for mature females and also attracts males and older nymphs, but 

to a lesser extent. It is produced from the ventral abdominal epidermis 

(Lucchi 1994). The principal ingredient of the mixture of compounds 

(obtained from extracts of male cuticle), isolated from bugs collected in 

southern France, is the sesquiterpene (z)-a-bisabolene trans epoxide, 

whereas the accompanying cis isomer is not attractive (BrŽzot et al. 1993). 

The pheromone blend differs between some Brazilian (which do not produce 

the cis compound) (Baker et al. 1987) and North American, Hawaiian and 

Japanese bug populations which do, but in differing ratios. Other related 

pentatomids (e.g. the North American Acrosternum hilare) emit mixtures 

containing distinctive ratios (but containing more cis isomers) of the same 

sesquiterpenes as in N. viridula (Aldrich et al. 1989). Males of the native 

Japanese Nezara antennata and Acrosternum aseadum produce speciesspecific 

pheromone blends based on the same compounds as Nezara 

viridula. However, whereas the trans/cis 1,2 epoxide ratio of N. antennata is 

within the range for most USA. N. viridula populations (3 to 4.4:1), the 

blend from Japanese N. viridula males is 0.82:1 to 1:1. The ratio for Italy 

was 2.16:1, for Brazil 2.28 to 4.67:1, and for Australia 3.90:1. The ratios for 

Acrosternum hilare, A. marginatum and A. pennsylvanicum are 6:100, 7:100 

and 94:100 respectively (Aldrich et al. 1989, 1993). However, the situation 

is not clearcut. Brezot et al. (1994) studied the proportions of cis and trans 

bisabolene epoxides in individuals of a southern France (SF) and a French 

West Indies (FWI) strain of N. viridula. The trans isomer composed 42 to 

82% of bisabolene epoxides in SF males and 74 to 94% of FWI males. 

Means differed significantly in spite of this inter-individual variation. Ryan 

et al. (1995) also found variability in the ratio of isomers within a single 

N. viridula population in Australia. 

It is interesting that the pheromone mixture from mature males in 

southeastern United States is also highly attractive to the tachinid parasitoid 

Trichopoda pennipes (Aldrich et al. 1987), which lays far more eggs on male 

than on female N. viridula. Trichopoda spp. and a group of related tachinids 

are native to the Americas, where they attack a small group of native 

pentatomid bugs. It is postulated that the chemical similarity of the 

Acrosternum and Nezara pheromones facilitated the adoption by the 

tachinids of N. viridula when it reached the Americas (see below). 

Furthermore, that the immigrant populations of N. viridula released both the 

trans and cis isomers and that parasitisation by the tachinids preferentially

4.12 Nezara viridula 209 

attracted to the cis isomer provided the major selection pressure leading to 

the present N. viridula populations having predominantly the trans isomer in 

their pheromone mix. Perhaps in parallel, in the 200 years or so of interaction 

between N. viridula and T. pennipes in tropical America, N. viridula has 

evolved a shorter pre-oviposition period and a longer developmental period 

than an Italian population (Hokkanen and Pimentel 1984; Aldrich et al. 

1989). At all events, at least two distinct pheromone strains of N. viridula 

can now be distinguished, based on the presence or absence in the volatile 

secretions of the cuticle of cis-(z)-a-bisobolene epoxide. Mature N. viridula 

males of one population from Brazil produce the trans, but not the cis isomer 

(Baker et al. 1987), whereas males from 2 other populations do in ratios of 

2.28:1 and 4.67:1 respectively (Aldrich et al. 1993), similar to males from 

southern USA which produce the trans and cis isomers in a 3:1 ratio (Aldrich 

et al. 1987) and those from Southern France in a 2:1 ratio (Baker et al. 1987). 

A quite different, but somewhat analogous situation, occurs with the 

hymenopteran Trissolcus basalis. A short chain unsaturated aldehyde, (E)- 

2-decenal present in the defensive scent produced in the adult N. viridula 

metathoracic gland (Gilby and Waterhouse 1965) is attractive to female 

T. basalis. A different compound, secreted on the eggs by the ovipositing 

female N. viridula, attracts female T. basalis to the egg raft (Mattiacci et al. 

1991, 1993). This compound is produced in the bug ovary and serves as an 

adhesive for attaching the eggs to the oviposition substrate. The adhesive 

and the material(s) responsible for the kairomone activity were partly 

soluble in water and completely in acetone and elicited recognition 

behaviour from T. basalis females when applied to glass beads. The 

adhesive appears to be a mucopolysaccharide-protein complex (Bin et al. 

1993). 


Early attempts at biological control were made chiefly in Australia and the 

Pacific, countries where the introduction of the bug was comparatively 

recent, but many other countries have been involved in more recent times. 

AUSTRALIA 

N. viridula was first reported in Australia in 1916 and soon became a 

widespread and serious pest. In 1933 the scelionid wasp Trissolcus basalis 

was introduced from Egypt into Western Australia (Table 4.12.2) where it 

readily became established and produced a great reduction in the pest status 

of the green vegetable bug. From Western Australia the parasitoid was 

distributed widely throughout south and southeastern Australia, making a 

considerable impact on the bug, except in cultivated areas in inland eastern 

Australia, where the cold winters affected its abundance (Wilson 1960).


Additional strains were later liberated of what, at the time, was believed to be 

T. basalis. These originated from the West Indies (1953), Italy (1956) and 

Pakistan (1961). However, doubt has been cast on the specific identity of the 

Italian material, although T. basalis is known to occur there. The Pakistan 

material, originally identified as T. basalis (but probably containing two 

species) has recently been shown (Clarke 1993a) to have consisted mainly of 

a new species Trissolcus crypticus. This bred readily in N. viridula eggs in 

the laboratory and, later, was widely distributed in Australia and Hawaii. 

However, T. crypticus has not been reported since in field collections. 

For many years up to the mid 1960s N. viridula was a very common and 

serious pest in the Canberra district, damaging tomatoes and beans in 

particular. This situation changed dramatically following the liberation of 

the material from Pakistan (presumably both T. basalis and T. crypticus) 

and, since then, N. viridula has become a very uncommon insect, appearing 

only in extremely limited numbers late in the season and only every few 

years. A great improvement also resulted about the same time in other 

subcoastal eastern cultivated areas, credited by Ratcliffe (1965) also to the 

liberation of material from Pakistan. 

However, since T. crypticus apparently did not become established its 

role, if any, in the changed situation is unclear. It is not known whether any 

cross mating with T. basalis might have occurred, which might have 

resulted in greater adaptability of T. basalis to the Canberra environment. 

The continuing very low abundance of N. viridula is possibly attributable to 

a heavy attack on its eggs by a resident population of T. basalis which is 

maintained on native pentatomid hosts, of which there are several (see later).

Table 4.12.2 Introductions for the biological control of Nezara viridula 


ANTIGUA 


Anastatus sp. 1961Ð62 Pakistan Ð Cock 1985 

Trichopoda pilipes 1949 

1955 

Florida 

Montserrat 

Ð 

Ð 

Cock 1985 

Cock 1985 

Xenoencyrtus hemipterus (= X. niger) 1963 Australia Ð Cock 1985 

ARGENTINA 

Trissolcus basalis 1981 Australia, Hawaii + Crouzel & Saini 1983; Porta & Crouzel 1984 

AUSTRALIA 

Bogosia antinorii 1958 Kenya _ Greathead 1971 

Telenomus chloropus 

(= T. nakagawai) 

1962 

1980 

1981 

Japan 

Japan 

Japan 

Ð 

Ð 

? 

Callan 1963 

Field 1984 

Field 1984; J. Turner pers. comm. 1984 

Trissolcus basalis 1933 Egypt + Kamal 1937; Wilson 1960 

1953 West Indies + Wilson 1960 

1956 Italy + Wilson 1960 

1961 Pakistan + Ratcliffe 1965 

1979Ð82 USA + Field 1984 

1979Ð82 Brazil + J. Turner pers. comm. 1984 

1979Ð82 South Africa + J. Turner pers. comm. 1984 

Trissolcus crypticus 1961 Pakistan Ð Clarke 1993a 

Trissolcus mitsukurii 1962 Japan + Callan 1963 

Trichopoda giacomelli Argentina ? Liljesthršm 1994 


Table 4.12.2 (contÕd) Introductions for the biological control of Nezara viridula 

Country and species Liberated From Result Reference 

Trichopoda pennipes 1941Ð43 

1949Ð50 

1952Ð53 

1980 

1980 

? 

Trichopoda pilipes 1952Ð54 

1980 

1980 

Ooencyrtus submetallicus 1953Ð57 

1962 

BRAZIL 

Florida 

Florida 

Florida 

Florida 

Florida 

Italy 

West Indies 

West Indies 

Hawaii 

Trinidad 

Trinidad 

Ð 

Ð 

Ð 

Ð 

Ð 

+ 

Ð 

Ð 

Ð 

Ð 

Ð 

Wilson 1960 

Wilson 1960 

Wilson 1960 

Michael 1981 

Michael 1981 

Giangiuliani et al. 1994; Colazza & Bin 1995 

Wilson 1960 

Michael 1981 

Michael 1981 

Wilson 1960 

CSIRO files 

Gryon japonicum Japan + Kishino & Teixeira 1994 

Gryon obesum USA + Correa & Moscardi 1995 

Ooencyrtus nezarae Japan Ð Kobayashi & Cosenza 1987 

Telenomus chloropus Japan Ð Kobayashi & Cosenza 1987 

Telenomus gifuensis Japan Ð Kobayashi & Cosenza 1987 

Trissolcus mitsukurii Japan + Kobayashi & Cosenza 1987; 

Kishino & Teixeira 1994, 

Trissolcus sp. Japan Ð Kobayashi & Cosenza 1987 

CALIFORNIA 

Trissolcus basalis 1987 

1987 

1987 

France 

Italy 

Spain 

+ 

+ 

+ 

Hoffman et al. 1991 






CHILE (EASTER ISLAND) 


Ectophasiopsis arcuata 1982, 1985/6 Chile + Ripa & Rojas 1989; Ripa et al. 1995 

Trissolcus basalis 1982 Chile Ð Ripa & Rojas 1989; Ripa et al. 1995 

COOK IS 

Trissolcus basalis 

FIJI 

1950 New Zealand ? Cumber 1953, Walker and Deitz 1979; 

A. Walker pers. comm. 1984 

Trissolcus basalis 1941 Australia + Lever 1941, 1943 


HAWAII 

1949 Florida ? OÕConnor 1950 

Trissolcus basalis 1962 Australia + Davis 1964, 1967 

Xenoencyrtus hemipterus 

(= X. niger) 

1962 Australia Ð Davis 1964, 1967 

Telenomus chloropus 1967 Japan Ð Davis & Chong 1968 

Telenomus sp. 1962 Australia Ð Davis 1964, 1967 

Trichopoda pilipes 1962 West Indies + Davis 1964, 1967 

Trichopoda pennipes 1962 Florida + Davis 1964, 1967 

Ooencyrtus submetallicus 1962 West Indies Ð Davis 1964, 1967 

Ooencyrtus trinidadensis 1962 West Indies Ð Davis 1964, 1967 

Trissolcus mitsukurii 1966 Japan Ð Davis & Krauss 1967 




ITALY 


Trichopoda pennipes 1984 or 

earlier 

1989 

Trissolcus basalis 1989 ? + Colazza & Bin 1995 

KIRIBATI 

? 

? 

+ 

+ 

G.K. Waite pers. comm. 

Gianguiliani & Farinelli 1995; Colazza et al. 1996a 


MONTSERRAT 

1979 Fiji + Anon. 1979b; Williams 1979; 

Dhamaraju pers. comm. 1985 


Trissolcus mitsukurii 

NEW ZEALAND 

1966 Japan Ð Cock 1985 

Ooencyrtus submetallicus West Indies Ð Jones 1988 

Trichopoda pennipes 1965Ð67 Florida Ð Cumber 1967; Clausen 1978 

Trissolcus basalis 1949 Australia + Cumber 1951 

Xenoencyrtus hemipterus 1962 Australia Ð Jones 1988 

NEW CALEDONIA 

Trissolcus basalis 1942Ð43 Fiji + Lever 1943 


Trissolcus basalis 1978 Australia + 

+ 

Anon. 1983 

Young 1982 

Trichopoda pennipes 1977 Hawaii Ð J.W. Ismay pers. comm. 1985 

Trichopoda pilipes 1980Ð81 Hawaii Ð Young 1982 




PITCAIRN IS 


Trissolcus basalis 1952 Fiji ? Dumbleton 1957 

POHNPEI 

Trissolcus basalis 1989 Hawaii + Esguerra et al. 1993 

AMERICAN SAMOA 

Trissolcus basalis 1953 Fiji ? Dumbleton 1957 

SAMOA 

Trissolcus basalis 1953 ? + Clausen 1978 

SOLOMON IS 

Trissolcus basalis 1940 Australia + CSIRO files 

Trichopoda pennipes 1940, 1949, 

1950 

SOUTH AFRICA 

Florida _ Dumbleton 1957 

OÕConnor 1950 

Trissolcus basalis 1955 Australia + Bedford 1964; Greathead 1971; Annecke & Moran 

1982; Bennett 1990 

Trichopoda pennipes 1986 

1994 

ST KITTS AND NEVIS 

Florida 

USA, Italy 

? 

? 

Bennett 1990 

Farinelli et al. 1994 


Trissolcus mitsukurii 1966 Japan Ð Cock 1985 

ST VINCENT 


TAIWAN 

Trissolcus basalis 1983 ? + Su & Tseng 1984 




TONGA 



USA (CALIFORNIA) 

1941 Australia + Dumbleton 1957 

Clausen 1978 


ZIMBABWE 

1992 eastern USA + Pickett et al. 1996 

Trissolcus basalis 1955 Australia ? Annecke & Moran 1982 



Doubt has been cast (Clarke 1990) both on the policy and effectiveness 

of introducing strains of T. basalis from different regions having differing 

environmental conditions. This practice has either led in Australia to 

effective biological control of N. viridula extending into additional 

environments or, alternatively, the initial genetic make-up of T. basalis has 

steadily undergone changes to allow progressive adaptation to new 

environments. It is possible that T. basalis consists of a complex of sibling 

species but, if not, it would be surprising if T. basalis has remained a 

homogenous species worldwide. Johnson (1985) found that American 

specimens of T. basalis showed much less morphological variation than 

those of Africa suggesting an African origin for the species. Furthermore, 

Handley (1975) reported that Australian T. basalis females would not mate 

with American males, although Powell and Shepherd (1982) found that 

reproductive isolation did not occur within any of 3 Australian strains 

examined; or between them and a strain from Florida. Nevertheless, the 

latter strain proved least fecund. Ferreira and Zamataro (1989) found no 

differences in reproductive capacity or longevity between an Australian and 

a Brazilian strain of T. basalis and Awan et al. (1989) concluded that Italian, 

French and Spanish populations consist of a single biotype, although several 

significant differences were observed in their biology. On the other hand, 

differences in courtship behaviour have been observed between different 

populations of T. basalis (Bin et al. 1988; Clarke and Walter 1992). 

However, it is not known whether any of these differences has any 

significance for biological control. 

Three other wasps have been introduced, Trissolcus mitsukurii (Japan 

1962), Telenomus chloropus (Japan 1962, Japan via USA 1980) and 

Ooencyrtus submetallicus (West Indies 1952Ð53). Only the former is 

believed to have become established but its impact has not been reported 

(Field 1984; J. Turner pers. comm. 1985). 

The only parasitoid reported from adult or nymphal N. viridula is the 

native tachinid Cylindromyia rufifemur (Cantrell 1984; Coombs and Khan 

1997). Three exotic species of parasitic tachinid fly have also been 

employed in attempts at biological control of Nezara viridula, Trichopoda 

pennipes from Florida, T. pilipes from the West Indies and Bogosia antinorii 

from Kenya. The Trichopoda species were introduced into Australia in the 

1940s and 1950s, but failed to become established (Wilson 1960). More 

recently the Trichopoda species were introduced into Western Australia 

from their native countries and also from Hawaii where they have been 

successfully established (Michael 1981). They have not become established 

in Australia. It is tempting to postulate that the pheromone blend secreted by 

Australian N. viridula does not attract Trichopoda pennipes and T. pilipes


females, whereas that of the Hawaiian population does. Whether or not this 

is true, it is clear that the responses by tachinid females to host pheromones 

introduces a considerable degree of host specificity to some particular 

populations of a host. 

The Argentinian Trichopoda giacomellii has been introduced to 

Australia for examination in quarantine for host specificity. Tests indicate 

that it has a limited host range involving only Nezara and its very close 

relations and it has now been approved for release in Australia (D.P.A. 

Sands pers. comm. 1997). 

At least three native wasps parasitise Nezara eggs, Telenomus sp., 

Xenoencyrtus hemipterus and ÔCoruna sp.Õ (certainly a misidentification of 

genus, Z. Boucek pers. comm. 1986), but these are of minor importance. 

The successful biological control of N. viridula in southern Australia has 

been repeated more recently in northwestern Australia where the green 

vegetable bug was first recorded in 1974. By 1976 populations were 

immense, for example over 33 nymphs and adults being recorded per square 

metre on a tomato crop and over 20 per head on badly affected sorghum. 

Since T. basalis was not present, it was introduced from southwestern 

Australia. Although more than 44 000 were released, initial establishment 

was poor. However, after 4 months, the situation improved dramatically, 

parasitisation was close to 100% and damage was reduced to sub-economic 

levels (Strickland 1979), although there are still periods of crop growth 

when the pest may be a problem (Michael 1981). 

T. basalis attacks, sometimes heavily, the egg masses of a range of 

pentatomid bugs. In southern Australia common pentatomid hosts are the 

horehound bug Agonoscelis rutila (Noble 1937; Clarke and Walter 1994) 

Cermatulus nasalis and Oechalia schellembergii (Awan 1989). In 

northwestern Australia alternative pentatomid hosts include Piezodorus 

hybneri and Oechalia schellenbergii, the egg masses of which suffer 

respectively 68% and 51% parasitisation. The coreid bug Riptortus serripes 

is also attacked, 38% of its egg masses being parasitised (Strickland 1979). 

Nezara viridula is under excellent biological control and is generally a 

very uncommon insect throughout southern Australia. However regular or 

occasional damage occurs in a subcoastal zone extending from south east 

Queensland (Titmarsh 1979) to north central New South Wales (e.g. 

Forrester 1979) and into northern Victoria (Clarke 1992a) and especially on 

soybeans. 

In an attempt to control these damaging populations, nine strains of 

T. basalis were introduced between 1979 and 1981, mass reared and 

released in southeastern Queensland. One strain came from each of South 

Carolina, Florida and Mississippi (USA), two from Brazil, two from South


Africa and two from northern Australia (Darwin and Kununurra). Although 

a slightly higher level of parasitisation has resulted, the problem has not been 

resolved (J. Turner, pers. comm. 1984; Clarke 1992a). It is notable that 

N. viridula is seldom a pest except in regions where soybean is a major crop. 

Turner (1983) showed that the rate of movement of T. basalis on soybean 

Glycine max was a third of that on cowpea Vigna unguiculata, mungbean 

V. radiata radiata, bean Phaseolus vulgaris, or sunflower Helianthus 

annuus. The proportion of N. viridula eggs parasitised on soybean was down 

to 25% of that on cowpea, mungbean and sunflower. Observations, since 

contested by Kelly (1987), suggested that the arrangement and height of the 

soybean leaf hairs, which are neither evenly spaced nor patterned, were 

responsible for interfering with the waspsÕ searching activities and this 

suggestion needs further investigation. Many soybean varieties have beeen 

selected for cicadellid resistance, which is directly correlated with the 

density, length and orientation of the leaf hairs (Broersma et al. 1972). 

Sesame Sesamum indicum leaves were repellent to the wasps, and those that 

did alight left immediately and engaged in vigorous grooming elsewhere 

(Turner 1983). Thus the nature of crops in an area can materially affect the 

success of biological control of the green vegetable bug. 

In spite of the foregoing, a claim has been made (Clarke 1990, 1992a,b, 

1993a,b, Clarke and Walter 1992), that, in Australia, Ôthere is little evidence 

to support claims of successful biological control of N. viridulaÕ (Clarke 

1993b). It is a mystery how such a view can be maintained in the light of the 

abundant evidence, available to its authors, from Western Australia, South 

Australia and coastal and southern New South Wales (Wilson 1960; Callan 

1963; Ratcliffe 1965; Strickland 1979; Michael 1981; Field 1984; 

Waterhouse and Norris 1987). Furthermore, until the 1970s, when 

increasing plantings of soybean, in particular, have provided highly suitable 

conditions for N. viridula populations to increase greatly in southern 

Queensland and northern New South Wales, there were even publications by 

entomologists in the Queensland Department of Primary Industries that, 

with the exception of the Darling Downs, Ôcontrol of the pest has become 

virtually unnecessary resulting from the introduction and establishment of a 

tiny parasite É Õ (Passlow and Waite 1971) and Trissolcus basalis Ôhas 

reduced the importance of Nezara viridula (L.) in coastal QueenslandÕ 

(Smith 1977). The highly effective control progressively achieved over a 

vast area of southern and Western Australia is in no way diminished in 

validity by the fact that N. viridula is, indeed, an important pest in a much 

smaller area extending from southeast Queensland, through central NSW to 

northern Victoria (Clarke 1992a). Although it is most commonly associated 

there with soybean (see later under Italy) it is also found on a range of other 

crops, including grain legumes, tomatoes and beans.


Clarke and Walter (1992) postulate that, because N. viridula oviposits 

only rarely during summer in southeast Queensland, T. basalis is largely 

without its preferred host for 60Ð90 days during which daily temperatures 

average more than 25¡C. This exceeds the average survival time of adult 

females at this temperature. Adult survival of T. basalis in summer is thus 

held to be the most likely factor limiting its populations. This postulate 

assumes that the egg masses of native pentatomid hosts of T. basalis are also 

in short supply over summer. It also appears not to apply to the far hotter but 

moister climate of the Ord Irrigation Area in far northern Western Australia, 

where N. viridula continues to be under generally excellent control. 

AFRICA 

N. viridula does not appear to be regarded as an important pest in northern 

Africa where its eggs are attacked by a number of Scelionidae, including 

Trissolcus aloysiisabaudiae, T. basalis, T. lepelleyi, T. maro, T. sipius and 

Telenomus seychellensis. Trissolcus basalis occurs mainly in coastal areas 

and the others are reported primarily from the eastern and central half of the 

continent. T. basalis was introduced to South Africa from Australia, New 

Zealand and USA, although there is some evidence that it may have already 

occurred there prior to these introductions (Giliomee 1958; Greathead 

1971). In Malawi eggs laid on macadamia were reported to experience an 

average of 74% parasitisation by Trissolcus maro and Telenomus 

seychellensis (Croix and Thindwa 1986). In Somalia Trissolcus 

alloysiisabaudiae is very abundant and may cause 100% parasitisation of 

N. viridula eggs on cotton (Paoli 1933). T. lepelleyi and T. sipius attack 

N. viridula eggs in East Africa, the latter being known only from Kenya. 

Psix striaticeps, which occurs in tropical Africa and India, has been bred 

from N. viridula eggs (Jones 1988). 

ARGENTINA 

N. viridula was first recorded in 1919. Since the native tachinid parasitoid 

Trichopoda giacomellii was unable to maintain populations at sufficiently 

low levels, three parasitoid wasps were introduced, of which the most 

effective is Trissolcus basalis (Crouzel and Saini 1983; Porta and Crouzel 

1984). 

In Buenos Aires Province, mortality of N. viridula eggs was found to be 

due mainly to parasitisation by T. basalis, that of 1st to 3rd instar nymphs to 

predation and that of adults (together with reduction in egg production) to 

parasitisation by Trichopoda giacomellii. Adult mortality and reduction in 

egg production was found to be density dependent. Three egg parasitoids 

were present, Trissolcus basalis (95% of total parasitisation), Telenomus 

mormideae and Telenomus sp.. Nymphal mortality was principally due to 

spiders and predatory bugs (Podisus sp.), although there was also some

BRAZIL 


parasitisation by Trichopoda giacomellii. Adverse climatic conditions 

(heavy rain) played a minor role in nymphal mortality (Liljesthršm and 

Bernstein 1990). Parasitisation of N. viridula eggs by T. basalis rose to a 

maximum of 90% in autumn although 33% of the parasitoids died, the 

majority (60%) in the pupal stage (Liljestršm and Camean 1992). 

In Rio Grande do Sul the main causes of mortality of N. viridula eggs laid 

throughout the season on soybean were infertility (2.7%: relatively 

constant), failure to hatch (14.1%: fluctuating), parasitisation (24%: 

relatively constant) and predation (17.3%: relatively constant) (Moreira and 

Becker 1986a). Three scelionid parasitoids were present, T. basalis, 

Trissolcus sp. and Telenomus mormideae. T. basalis killed a greater number 

of eggs and attacked a larger number of egg rafts than the other species 

(Moreira and Becker 1986b). A complex of polyphagous predators did not 

discriminate between parasitised and unparasitised eggs and were 

responsible for 25.5% mortality. The predators were responsible for 17.3% 

mortality of N. viridula and 34% of T. basalis (Moreira and Becker 1986c). 

Predation on host eggs was the main cause of mortality of T. basalis in the 

pre-emergence period (Moreira and Becker 1987). 

The tachinid fly Trichopoda giacomellii (= Eutrichopodopsis nitens) is 

the most important parasitoid of N. viridula in northern Paran‡ State. The 

level of parasitoid attack varies according to the plant on which the host is 

feeding and is highest when soybean is not available (Panizzi 1989). 

Although T. basalis (introduced) and Telenomus mormideae (native) were 

already present on the Cerrados area, Kobayashi and Cosenza (1987) 

introduced from Japan 5 additional species. In order of decreasing efficacy 

in parasitisation and adult emergence these were Trissolcus mitsukurii, 

Ooencyrtus nezarae, Telenomus chloropus, Telenomus gifuensis and 

Trissolcus sp.. Of the introduced species, T. mitsukurii parasitised eggs of all 

major pentatomid species throughout the year and also survived the dry 

winter season. In addition, it was the dominant competitor on egg masses. 

However, Bennett (1990) reports, more recently that it is not definite that 

permanent establishment has been achieved. When compared with 

T. basalis, the latter parasitised about 90% of exposed eggs with 60% adult 

emergence, whereas T. mitsukurii achieved about 70% parasitisation and 

40% emergence (Kobayashi and Cosenza (1987). In northern Brazil, 

Ferreira (1986) reported 40% parasitisation of N. viridula eggs by T. basalis 

and that Telenomus mormideae was also abundant. 

In the Federal District of Brazil the egg parasitoids Trissolcus mitsukurii 

and Gryon japonicum, introduced from Japan, gave good levels of 

parasitisation (Kishino and Teixeira 1994). In Parana State T. basalis,


Telenomus podisi and Gryon obesum parasitised up to 60% of Nezara eggs 

on soybean (Correa and Moscardi 1995). 

In southern Brazil, T. basalis is the main parasitoid, attacking between 

97.5% and 100% of N. viridula eggs laid on soybean. In 1988, 36.6% of the 

egg masses were attacked and 10.3% in 1989, with 21.8% and 6.3% of the 

individual eggs being parasitised respectively (Foerster and QueirÏz 1990). 

COOK IS 

T. basalis was introduced to Mangaia in 1950, but is not known to have 

become established (Cumber 1953; Walker and Deitz 1979) and this must be 

assumed not to have occurred. 

EASTER IS 

Control of N. viridula was achieved by the establishment of the tachinid 

Ectophasiopsis arcuata from mainland Chile, so that it is now difficult to 

find a bug. T. basalis was also introduced but was not recovered (Ripa and 

Rojas 1989; Ripa et al. 1992). 

FIJI 

N. viridula was first recorded in 1939 and Trissolcus basalis was introduced 

from Australia in 1941 (Lever 1941). Success was immediate and good 

control resulted (OÕConnor 1950). Large populations of N. viridula are 

reported to develop sometimes on cowpeas, but the insect is not troublesome 

on other legumes (Swaine 1971). 

Trichopoda pennipes was introduced from Florida in 1949, but its 

establishment is not recorded (OÕConnor 1950). 

HAWAII 

N. viridula was first recorded in 1961 and the wasps Trissolcus basalis, 

Xenoencyrtus hemipterus (= X. niger) and Telenomus sp. (all egg 

parasitoids) from Australia were released in 1962. Other importations in 

1962 were the tachinid fly, Trichopoda pilipes, which parasities last instar 

nymphs and adults, and two egg parasitic wasps Ooencyrtus submetallicus 

and O. trinidadensis from the West Indies. In 1963 Trichopoda pennipes 

was imported from Florida. Of these parasites, Trissolcus basalis, 

Trichopoda pennipes and T. pilipes became established (Davis and Krauss 

1964; Davis 1964, 1967; Croix and Thindwa 1967; Clausen 1978). Nezara 

populations declined steadily to sub-economic levels, with only sporadic 

outbreaks, and the species is generally under effective biological control 

(C.J. Davis, pers. comm. 1985). Average parasitisation by Trissolcus basalis 

ranged up to about 95% and by Trichopoda pilipes up to 86%. Trichopoda 

pupae are occasionally parasitised by the encyrtid Exoristobia philippensis 

(Davis 1964). 

More recently (1990Ð91) egg rafts of N. viridula placed in weeds at the 

border of macadamia nut plantations had significantly higher rates of


parasitisation (49.9%) than rafts placed in the canopy of macadamia trees 

(14.7%). Predators were more effective at locating rafts placed in trees than 

in weeds and were always more efficient than T. basalis, regardless of their 

location. The egg parasitoid, Anastatus sp. was equally inefficient in both 

habitats. In 1990 only 1.2% of the eggs in the trees were parasitised and 8.6% 

in the weeds. During the same period, predators destroyed 26.0% and 14.5% 

in trees and weeds respectively. 

In 1991 parasitisation of eggs dropped to 0.2% and 1.7% in trees and 

weeds, whereas predation increased to 47.7% and 36.9% respectively. 

Doubt was, therefore, cast upon T. basalis having a prominent role in 

biological control of N. viridula in Hawaii (Jones 1995). Although this 

conclusion appears to follow in the macadamia agroecosystem studied, it 

would be of interest to know whether it applies also to other susceptible 

crops. Predation was attributed mainly to ants, including Pheidole 

megacephala. 

INDIA 

Singh (1973) reported no parasitoids in life-table studies of N. viridula on 

soybeans. 

INDONESIA 

Partial life tables showed on soybeans in Northern Sumatra that mortality of 

N. viridula until the late 1st instar was 50 to 87%, of which 18 to 85% 

occurred during the egg stage and was caused mainly by predators. Only 2 to 

26% of the eggs were parasitised. The main predators were two species of 

ants (Solenopsis geminata and Dolichoderus sp. a staphylinid beetle 

(Paederus sp.) and several crickets, although other egg predators belonging 

to the families Tettigoniidae, Lygaeidae and Anthocoridae were also 

observed feeding on the eggs. Trissolcus basalis parasitised the eggs but no 

evidence was obtained of the presence of tachinid parasitoids that attack late 

nymphs and adults (van den Berg et al. 1995). 

ITALY 

Before production of soybeans began in Italy in 1981 N. viridula was only 

important occasionally. Crops attacked included tomatoes and legumes. The 

increasing production (over a decade more than a thousand fold increase in 

area planted to soybeans) filled a temporal and food gap for N. viridula and 

other pentatomids (Colazza and Bin 1990). The second generation of 

N. viridula now migrates each summer into soybeans at the beginning of 

development of pods, which then provide the main food for reproduction 

and larval development. Abundance of N. viridula increased steadily in the 

eighties throughout northern and central Italy to a level at which it became a 

key pest. Three parasitoids were recorded from egg rafts, Anastatus 

bifasciatus, Ooencyrtus sp. and Trissolcus basalis. However the first 2


JAPAN 

species were never bred from egg rafts collected from soybeans. 

Approximately 20% of egg rafts were parasitised in 1986 and 1987, 

increasing to 50% in 1988 to 1992. Efficiency of parasitisation (% eggs 

parasitised divided by the number of egg masses discovered) was 65% in 

1986, but rose to 92% in 1988 and 1990. 

The early larval instars were generally free from parasitoid attack 

although two tachinids were occasionally recorded, the native Ectophasia 

crassipennis (5 to 10% parasitisation) and the accidentally introduced 

Trichopoda pennipes (2 to 15% parasitisation). 

T. basalis was also recovered from the eggs of 2 other pentatomid bugs 

present at low levels in soybean fields, Carpocoris mediterraneus and 

Piezodorus lituratus (Colazza and Bin 1995). 

The widespread Telenomus chloropus (= T. nakagawi) and also Trissolcus 

(= Asolcus) mitsukurii which is known only from Japan are the two most 

important parasitoids of N. viridula and have been studied intensively in 

Fukuoka. The host-finding ability of T. chloropus is superior to that of 

T. mitsukurii and it parasitises egg masses more rapidly and more 

efficiently. Females lay about 100 eggs (which is 1.6 times that of 

T. mitsukurii and live 11 days, or 3 days longer than T. mitsukurii (Nakasuji 

et al. 1966), although the latter may have at least 11 generations a year, 

whereas the former has some 9 generations (Hokyo et al. 1966b). Hokyo et 

al. (1966a) have shown experimentally that the two species do not 

discriminate between each otherÕs parasitised and unparasitised eggs, with 

T. mitsukurii larvae usually being successful in competition with 

T.chloropus larvae (Hokyo et al. 1966a). Furthermore, female T. mitsukurii 

often bite and kill female T. chloropus when they meet on the egg mass 

(Hokyo and Kiritani 1966). It follows that the effectiveness of T. chloropus 

is reduced by the presence of T. mitsukurii (Nakasuji et al. 1966), so it would 

be undesirable, in a biological control program, to introduce the latter along 

with the former. However, it should be borne in mind that T. mitsukurii is 

more abundant than T. chloropus in the southern coastal district of Fukuoka, 

whereas the reverse is true for the northern mountainous districts. The 

combined mortality caused by the two species amounted to 60 to 90% of the 

first spring generation N. viridula eggs. 

Three minor parasitoids have been bred from Nezara eggs in Fukuoka 

and several others are present elsewhere. The first, Telenomus gifuensis, is 

an effective parasitoid of Scotinophara lurida eggs, and also attacks a range 

of other pentatomids (Hidaka 1958; Hokyo et al. 1966b). The second, 

Ooencyrtus nezarae, the smallest of the three, is known from Nezara 

viridula, N. antennata and Anacanthocoris concoloratus. The third,


Anastatus japonicus, is known as an egg parasitoid of the gypsy moth 

Lymantria dispar and other Lepidoptera (Hokyo et al. 1966b). 

KIRIBATI 

N. viridula was a pest on the islands of Betio and Tarawa in the 1970s. 

T. basalis was released in 1978 and, since 1984, this pest has not been 

recorded from Tarawa (E. Dharmaraju pers. comm. 1985). 

NEW CALEDONIA 

Trissolcus basalis was introduced in 1942Ð43 and became established 

(Lever 1943; Clausen 1978). 

NEW ZEALAND 

N. viridula was first recorded in 1944 and soon became a serious pest of 

many crops. T. basalis was introduced from Australia in 1949 and rapidly 

became widely established. There followed a gradual decline in the severity 

of plant damage and, although populations continued to fluctuate seasonally, 

the situation became satisfactory (Cumber 1949, 1951, 1953, 1964). Over 

this period T. basalis extended its host range to other pentatomid bugs (e.g. 

Cuspicona simplex and Glaucias amyoti), thereby providing a source of 

parasitoids to attack any eggs of Nezara that became available. Adaptation 

of T. basalis to Nezara under New Zealand conditions may also have been 

responsible for its improved performance (Cumber 1964). 

Trichopoda pennipes, originally from Florida, was obtained from 

Hawaii in 1965 and released over the next 3 summers. Evidence of a 

generation in the field was obtained in May 1967, but the fly did not become 

established. In addition to Nezara viridula, eggs were deposited on adults of 

other pentatomids Antestia orbona, Cermatulus nasalis, Cuspicona simplex, 

Glaucias amyoti and Dictyotus caenosus, but a parasitoid was reared only 

from G. amyoti whose nymphs are readily parasitised (Cumber 1967). 


N. viridula is a serious pest in the Markham Valley where T. basalis is 

present, but generally results in less than 30% parasitisation. In 1978 a strain 

of this parasite from Western Australia was released, but the level of 

parasitisation did not increase (Young 1982). In Wau T. basalis and another 

scelionid egg parasite are generally effective, although N. viridula 

occasionally increases to pest proportions (Gagne 1979). The tachinid 

parasites Trichopoda pennipes and T. pilipes were introduced from Hawaii 

but failed to become established (J.W. Ismay pers. comm. 1985). 

T. giacomelli is being considered for release. 

PHILLIPINES 

Three decades ago it was reported that N. viridula was not a pest, apparently 

being controlled by a native egg parasitoid, Ooencyrtus sp. (Cendana, in 

Davis 1967) and it is interesting that Nezara was reported as present, but 

unimportant, in 1993 (Waterhouse 1993b). This may possibly be correlated


with the fact that soybean production in the Philippines is rather limited 

Ñmuch lower than in many other Southeast Asian countries, such as 

Thailand, Indonesia and Vietnam. Two egg parasitoids Telenomus comperei 

and T. pacificus have been reported from N. viridula eggs laid on 

groundnuts. Both species parasitised 100% of offered eggs in 24 hours and 

adults emerged after 12 to 14 days. Adults lived up to 32 days when fed 

honey (Cadapan and Alba 1987). 

POHNPEI 

N. viridula became a major pest in the early 1990s on several islands in the 

Federated States of Micronesia. Following the introduction of T. basalis 

from Hawaii, the green vegetable bug population has become so low that it is 

rarely seen now on vegetables in Pohnpei (Esguerra et al. 1993; Suta and 

Esguerra 1993). 

SAMOA 

Trissolcus basalis was introduced in 1953 and became established (Clausen 

1978). 

SOLOMON IS 

Trissolcus basalis was introduced from Australia in 1940 against the 

coconut spathe bug Axiagastus campbelli. It is said to be established 

(CSIRO files) and a Trissolcus sp. has been recorded from pentatomid eggs 

(N. viridula or Plautia brunneipennis) on beans (R. Macfarlane pers. comm. 

1985). 

Trichopoda pennipes was introduced from Florida via Fiji in 1950 in 

order to control the coconut bug Amblypelta cocophaga and other 

phytophagous bugs (OÕConnor 1950), but it has not been collected since. 

SOUTH AFRICA 

Trichopoda pennipes was introduced from USA and Italy and liberated in 

1994 (Farinelli et al. 1994; van den Berg et al. 1994) but there is no 

information on establishment. T. giacomelli has also been imported for 

study (D.P.A. Sands pers. comm. 1997). 

TAIWAN 

T. basalis was introduced in 1983 and, two months after release, 

parasitisation rates of 90% and 60% respectively of N. viridula eggs at two 

sites was reported (Su and Tseng 1984).

THAILAND 

TONGA 

USA 

VANUATU 


The most abundant egg parasitoid is Gryon fulviventris, which exists as a 

number of biotypes. Under a series of synonyms (Dissolcus fulviventris, 

Hadronotus fulviventris, H. antestiae and Gryon antestiae) it is known from 

Africa, India, Thailand, southern USSR and Malaysia. It parasitises the eggs 

of many species of Pentatomidae, Scutelleridae and Coreidae, but was 

reported by Jones (1988) for the first time in N. viridula eggs in Thailand, 

where it also breeds in the eggs of Piezodorus hybneri. In Africa larvae 

develop in Nezara viridula eggs, but adults do not emerge successfully. 

Other egg parasitoids are Telenomus chloropus, Ooencyrtus nezarae, 

Anastatus sp. (Jones et al. 1983b; Jones 1988), Telenomus sp. and Trissolcus 

basalis (Napompeth 1990). 

Trissolcus basalis was imported in 1941 and became established (Clausen 

1978). 

The influence of the host plant on which egg rafts of N. viridula are laid on 

the level of both parasitisation and predation was investigated in North 

Carolina by Shepard et al. (1994). Parasitisation was higher than predation 

on eggs on tomato and about equal in okra, soybean and cowpea. In one year, 

predation was higher than parasitisation in soybean towards the end of the 

growing season, but parasitisation was higher early in the season in okra and 

cowpea. Parasitisation of egg masses in wild radish reached a peak of nearly 

100% during spring and declined to about 30% in autumn. The major 

parasitoid from all crops was Trissolcus basalis, although Ooencyrtus 

submetallicus occured in low numbers. The conclusion was reached that 

both parasioids and predators may play an important role in regulating 

populations of N. viridula and that their combined action often resulted in 

the attack of 100% of egg masses in some crops. 

N. viridula occurs in Vila where its eggs are heavily parasitised by a wasp 

Trissolcus sp. (not T. basalis) (R. Weller pers. comm. 1986).


Biology of the major species 

Scelionidae: Hymenoptera 

This is the most important family of hymenopterous parasitoids emerging 

from the eggs of N. viridula and is dealt with, amongst others, by Nixon 

(1935, 1936, 1937, 1966). 


Kamal (1937) was the first of many to make a detailed study of the impact of 

the egg parasite Trissolcus basalis on the abundance of Nezara viridula: no 

control measures are needed in Egypt. His work on the biology of the wasp 

has been supplemented by studies by many later workers (e.g. Wilson 1961; 

Cumber 1964; Powell and Shepherd 1982; Correa and Moscardi 1993, 1994; 

Awadalla 1996; Colazza et al. 1996b). The minute female of T. basalis 

oviposits in the side of the bug egg, after which she marks the egg by rubbing 

an abdominal secretion over it with the ovipositor, as a deterrent to other 

females from laying in the same egg. The length of the life cycle ranges from 

9 to 24 days depending on temperature, the entire egg, larval and pupal 

stages being passed inside the same eggshell. The adult wasps chew their 

way out through the lid of the eggshell, males usually emerging first and 

disputing with one another for possession of the egg batch and thereby the 

right to fertilise the later-emerging females. In hot weather the females live 4 

to 15 days and have considerable dispersive powers, as shown by the 

rapidity with which they spread through newly colonised areas. The adult 

wasps overwinter among leaves and litter. 

As indicated earlier, there is good evidence for the existence of several 

different strains of T. basalis. Experimental work using strains from three 

widely separated regions of Australia and from Florida showed that they 

were not reproductively isolated, that the Australian strains had a higher 

fecundity, but that adults of the Florida strain lived longer (Powell and 

Shepherd 1982). 

T. basalis is frequently recorded from several other pentatomids, but has 

a special preference for N. viridula (Jones 1988). It appears to be most 

effective in coastal and subcoastal areas and has been established in 

Argentina, Australia, Fiji, Hawaii, Kiribati, Papua New Guinea, New 

Caledonia, Samoa, Solomon Is, South Africa and Tonga and possibly in 

several other countries (Table 4.12.2). 

In Australia two species of the pteromalid Acroclissodes are parasitic on 

T. basalis (Clarke and Seymour 1992).


Gryon sp. 

Some aspects of mating and reproduction of Gryon sp. in India have been 

investigated by Velayudhan and Senrayan (1989). 

Trissolcus mitsukurii 

This important egg parasitoid of Nezara in Japan also attacks the eggs of 

several other pentatomids (Kishino and Teixeira 1994) , preferring species 

that deposit their eggs in small masses. It is bisexual and the first egg 

deposited by a mated female always produces a male. Both sexes have 

aggressive behaviour and females drive Telenomus chloropus females off a 

pentatomid egg mass (Hokyo et al. 1966b). The fecundity and longevity of 

T. mitsukurii were found to be less than those of T. basalis in laboratory 

trials at 26¡C and 65% RH. T. basalis parasitised 82.2% of eggs on the 

second day of adult life, whereas T. mitsukurii parasitised only 51.3%. On 

average, the former laid 250 eggs and the latter 80, and the longevity of 

T. basalis was 80.1 days and of T. mitsukurii 42.6 days (Ferreira and 

Zamataro 1989). 

Too little is known about the dozen other Trissolcus species in Table 

4.12.1 to form an opinion of their value in biological control. 

Telenomus chloropus (= T. nakagawai) 

This is one of the most important egg parasitoids of N. viridula in Japan and 

also attacks eggs of N. antennata. Females are parthenogenic, lay about 100 

eggs and live for 11 days. They have a high searching ability and can 

parasitise all egss in a raft (Nakasuji et al. 1966). The species is oligophagous 

and prefers large egg rafts of pentatomids, such as those of N. viridula or 

N. antennata, to smaller egg masses of many other pentatomids. When 

introduced to the laboratory in Louisiana, USA. females lived for 8 days at 

24¡C, laid on average 60 eggs and developed from oviposition to emergence 

in 18 days. When reared from eggs of N. viridula which had been reared on 

resistant soybean its fecundity was half that of parasitoids reared on 

susceptible soybean and its mortality within Nezara eggs was higher (Orr et 

al. 1985b). 

Although morphological differences have not been found, it appears that 

T. chloropus exists as a series of biotypes. It is a widespread polyphagous 

parasitoid of pentatomid eggs throughout the Palaearctic Region, but it is 

recorded from Nezara only in Japan, Korea and Thailand. The Japanese 

biotype rarely produces males, although males occur elsewhere (Jones 

1988). T. chloropus from Japan has been released in Australia (Callan 1963; 

Field 1984), Brazil (Kobayashi and Cosenza 1987) Hawaii (Davis and 

Chong 1968) and the USA (Jones 1988), but has not become established, 

possibly due to its requirement for high humidity (85% RH or higher) for 

successful emergence (Orr et al. 1985a).


Telenomus cyrus 

This parasitoid is known from Indonesia, the Philippines and Taiwan. In 

Taiwan it parasitises up to 19% of eggs in soybean and is the most important 

egg parasitoid of N. viridula in soybean, rice and jute (Taiwan Agricultural 

Research Institute 1984). 

Encyrtidae: Hymenoptera 

Species of Ooencyrtus have been recorded from N. viridula eggs from many 

parts of the world (Table 4.12.1), but are never a major component of the 

suite of parasitoids. 

Ooencyrtus submetallicus 

This species ranges from Florida through to West Indies, Brazil and 

Argentina (Jones 1988). It was found to be inferior to T. basalis in host 

location and dispersal in soybeans in Louisiana (Lee 1979). It was 

introduced to Australia, Hawaii and New Zealand, but did not become 

established (Wilson 1960; Davis and Krauss 1963; Davis 1967). 

Tachinidae: Diptera 

These are clearly important parasitoids of adult N. viridula in the Americas 

and the Ethiopian region. Outside of Japan, where one species attacking the 

green vegetable bug is known, there appear to be no records of tachinids 

regularly attacking N. viridula in Eastern Asia. 

Trichopoda spp. 


T. pennipes in North America is a complex of biotypes or sibling species. In 

the east its native hosts are the squash bug Anasa tristis, other coreids and 

several pentatomids (Arnaud 1978). In the southeast it occurs regularly in 

the native pentatomid Acrosternum hilare, but seldom in other pentatomids 

(Jones 1988). In California it does not oviposit on A. tristis, but in the field 

breeds in a pyrrhocorid and a largid bug (Sabrosky 1955; Dietrick and van 

den Bosch 1957). Salles (1991, 1993) has studied T. pennipes in Florida. 

The female Trichopoda pennipes lays eggs singly on the cuticle, mainly 

of the undersurface of fourth and fifth instar nymphs and adult bugs. The 

eggs hatch in 3 to 4 days and the young larvae bore directly into the host, tap 

the respiratory system of the bug for air and feed on the body fluids and 

internal organs of the host. When fully fed (16 days), the third instar larvae 

forces its way out through an intersegmental membrane of the host abdomen 

and pupates in the soil. After about 14 days the adult fly emerges. Up to 232 

eggs are laid by a female and, unlike Trissolcus basalis, there is often great 

wastage, many eggs (up to 237) being laid by several females on one adult 

bug, although only one parasitoid larva survives (Shahjahan 1968). The


reproductive organs of the host bug may or may not be aborted by the 

feeding of the parasite, and it ultimately dies from mechanical injury caused 

by the emerging larva. The parasitoid overwinters as a second instar larva 

inside the hibernating adult bug (Beard 1940; Clausen 1978). Rearing 

methods are discussed by Gianguiliani and Farinelli (1995). Male 

N. viridula receive more parasitoid eggs than females and are more heavily 

parasitised (Mitchell and Mau 1971; Todd and Lewis 1976). 

In Georgia, USA, female Nezara parasitised by T. pennipes live about 

half as long as normal females and lay about a quarter the number of fertile 

eggs (Harris and Todd 1980). 

Trichopoda pilipes 

In the West Indies, instead of T. pennipes, the closely related T. pilipes 

(sometimes regarded as a subspecies) occurs. Both species have been 

established in Hawaii and T. pilipes is the more important (Davis 1967). 

Unsuccessful attempts have been made to establish one or both of these 

species in Australia, Fiji, South Africa and a number of other places, 

although T. pennipes has been established in Italy (Table 4.12.2). 

Trichopoda giacomellii 

Parasitisation levels by T. giacomellii of N. viridula in Argentina were 

45.3% on sorghum, 42.1% on flax, 29.9% on wheat and 27.9% on soybean. 

Levels on males were higher than on females, except on soybeans, where 

there was no significant difference (La Porta 1990). Liljesthršm (1985, 

1995) observed that the highest densities of parasitoids and the highest rate 

of parasitisation occurred in areas with the highest densities of N. viridula. 

Many fly eggs are laid on some individual hosts and few or none on 

others. More eggs are deposited on adult N. viridula than on 4th or 5th instar 

nymphs and more on adult males than on females. Some bugs are attacked 

sufficiently late in their development that they are able to produce at least 

one normal egg batch which has unaffected egg viability. In one study less 

than 7% parasitised nymphs died in the 5th instar. This resulted in sufficient 

eggs being laid to enable N. viridula to persist in the environment 

(Liljesthršm 1992, 1993a,b). In the laboratory at 26¡C, 70% RH and a 16hour 

day, the egg, larval and pupal stages lasted 2.8, 33.0 and 13.3 days 

respectively and females laid an average of 29 eggs (La Porta 1987). 

On hatching, young larvae penetrate the host cuticle and, on moulting to 

the 2nd instar, attach their posterior spiracles to one of the hostÕs tracheal 

trunks. The fully grown 3rd instar larva emerges from the host to pupate in 

the soil. Most hosts die shortly after the parasitoid larva has left. 

In one large field sample the maximum number of living parasitoid 

larvae found per host was 2 (4% of hosts), whereas only 1 living parasitoid


larva was found in 66% of hosts. Small, dead, damaged parasitoid larvae in 

some hosts provided evidence that there was competition for survival 

(Liljesthršm 1993b). 

Because moulting led to loss of unhatched eggs with the discarded 

cuticle, eggs laid on nymphs less frequently led to successful parasitisation 

than eggs laid on adults. With 1 parasitoid egg per adult, success was greater 

on males than on females whereas, when more than 4 eggs were present, the 

success rate was higher with females (Liljesthršm 1991). T. giacomellii 

parasitised 100% of N. viridula adults for 3 consecutive generations in an 

uncultivated area near Buenos Aires (Liljesthršm 1981) and it was 

concluded that T. giacomellii could regulate the population of N. viridula 

(Liljesthršm and Bernstein 1990). 

T. giacomellii (at times referred to incorrectly as Eutrichopodopsis 

nitens) is also an important parasitoid of N. viridula in Brazil. When 

parasitisation occurred in nymphs or newly moulted adults, adults did not 

reproduce and longevity was greatly reduced. Female N. viridula, 

parasitised on the 7th day of the adult stage, had their fecundity reduced by 

58%, but neither egg fertility nor size was affected (Ferreira et al. 1991). 

Parasitisation by T. giacomellii collected in the field in Brazil from soybean 

and other crops ranged from 27.1% to 52.7%. More parasitised eggs were 

found on males than on females and most eggs were on the thorax (Ferreira 

1984). High rates of parasitisation of N. viridula by T. giacomellii were 

observed on the weed Leonurus sibericus, but populations transferred to 

nearby soybean when this entered the reproductive phase. On the other hand, 

bugs living on castor, Ricinus communis, stayed on this plant all year round. 

This weed is of low nutritional value to them, but on it they are less liable to 

attack by the tachinid (Panizzi 1989). 

A recent laboratory study of the reproductive attributes of T. giacomelli 

determined the influence of adult food availability and body size on 

fecundity, and longevity, both relevant to any introduction program 

(Coombs 1997). 

Little is known about other Trichopoda species attacking pentatomid 

bugs: T. lanipes in Florida (Drake 1920), Trichopoda sp. in Uruguay (Guido 

and Ruffinelli 1956) and possibly other species in Brazil (Jones 1988). 

Bogosia antinorii 

This species is widespread in eastern and southern Africa and is known only 

from N. viridula (van Emden 1945; Barraclough 1985). There was 

apparently an unsuccessful attempt to establish it in Australia from material 

from Kenya (Greathead 1971).


Ectophasiopsis arctuata 

This tachinid is common on N. viridula adults in Chile. Following its 

introduction to Easter Is it brought this bug under successful biological 

control (Ripa and Rojas 1989). 

Gymnosoma rotundata 

This tachinid parasitises N. viridula in Japan where up to 5% parasitisation is 

recorded (Kiritani et al. 1963). It is widespread in Palaearctic regions and 

attacks many hosts, including Nezara antennata, in Japan and Korea. 

Gymnosoma clavata has been recorded once from N. viridula in Europe 

(Herting 1960). 

One other tachinid has been reported once from N. viridula, 

Cylindromyia rufifemur from Australia (Cantrell 1984). 

Comments 

Although predators are undoubtedly important natural enemies of 

N. viridula, particularly of its early stages, most are generalists which are 

unlikely to be approved nowadays by quarantine authorities for introduction 

as biological control agents. This situation is likely to be partly offset by the 

fact that most countries possess a suite of generalist predators, some of 

which are likely to attack N. viridula. 

The species of egg parasitoid most closely associated with N. viridula 

are concentrated in Africa and Japan. Elsewhere its eggs are parasitised by 

introduced species or by native species that have expanded their activities 

from native bugs. Indeed, it seems likely that the complex of Japanese 

parasitoids has probably expanded its host range from the oriental stink bug 

N. antennata of Japanese origin, just as the complex of Central and South 

American tachinids has clearly expanded to N. viridula adults from adults of 

native bugs. 

Successful biological control of N. viridula has been achieved in many 

countries to which it has spread this century. These include a vast area (but 

not all) of Australia, also New Zealand, Hawaii and several other Pacific 

islands. The prospects are good for reducing its pest status in many other 

areas where effective parasitoids are not yet present. This might involve the 

introduction of additional species or strains of Trissolcus. In addition several 

of the many other parasitoids known to attack eggs (species in the genera 

Telenomus and Gryon) are well worth considering. In the Ethiopian region a 

tachinid (Bogosia antinorii) is an important parasitoid of adults and large 

nymphs and in the Americas there are at least 4 tachinid species worthy of 

consideration: Trichopoda pennipes: USA; T. pilipes: West Indies;


T. giacomelli Argentina; and Ectophasiopsis arcuata: Chile. These 4 are 

reported to be more abundant now on N. viridula than on the native bugs 

they parasitised before the arrival of N. viridula (Jones 1988). A problem in 

their effective use is that location of N. viridula hosts is dependent, in some 

species at least, upon the secretion by N. viridula of a specific attractive 

blend of chemicals. Some biotypes of N. viridula that do not produce the 

appropriate blend largely escape oviposition. There are no records of 

tachinids regularly attacking N. viridula in the East Asian Region, except for 

the widespread, polyphagous Gymnosoma rotundata, which was found to 

cause up to 5% parasitisation of N. viridula in Japan and also to attack 

N. antennata in Japan and Korea (Kiritani et al. 1963). 

The main areas where N. viridula continues to be an economically 

important pest in spite of attempts to use natural enemies including 

T. basalis appear, with the exception of certain crops such as macadamia and 

pecan nuts, to be associated with extensive plantings of soybeans. A detailed 

re-examination is required of the behaviour of T. basalis (and perhaps other 

egg parasitoids) in relation to ability to parasitise N. viridula egg masses laid 

on soybean. If it is demonstrated that certain physical or chemical 

characteristics of soybeans are responsible for poorer than usual 

performance, serious consideration should be given to the selection of 

varieties that have minimal adverse effects on the parasitoids. This, of 

course, is different from selecting soybean cultivars that are resistant to 

N. viridula, some of which are known (e.g. Kester et al. 1984; Bowers 1990). 

In this context it is relevant that the biology of Telenomus chloropus, an 

egg parasite introduced into southern USA in 1982 from Japan, was studied 

on eggs of N. viridula that had been reared on the stink bug-resistant 

soybean, PI 717444, or on the susceptible cultivar, Davis. Time of 

development of the parasite did not differ significantly in eggs from either 

source, but success of emergence was lower from eggs laid on resistant 

soybean and fecundity of those that did emerge was about half of that of 

individuals reared from eggs laid on Davis. The authors (Orr et al. 1985b) 

point out that, with a marked reduction in emergence and fecundity, 

combined with decreased host availability, there is the potential for 

reduction or elimination of resident parasite populations in fields of resistant 

soybeans. 

Comparatively little is known of the range of Nezara parasitoids in its 

centre of origin, namely the Ethiopian region, although at least 6 Scelionidae 

including T. basalis have been recorded, together with the apparentlyspecific, 

widespread tachinid Bogosia antinorii, whose effectiveness 

deserves study. It is probable that a thorough investigation in the Ethiopian 

region would disclose an additional range of potentially valuable species.

4.13 Ophiomyia phaseoli 

India 

Myanmar 

+ 

20° 

Laos 

++ 

0° 

20° 

China 

P 

Thailand 

+ 

Cambodia 

Vietnam 

++ 

P 

++ Brunei 

Malaysia 

+ 

Singapore 

+++ 

Indonesia 

Taiwan 

P 

++ 

Philippines 

Australia 

Papua 

New Guinea 

++ 

235 

It appears that the bean fly Ophiomyia phaseoli originated in Asia. Its most effective 

natural enemy, the braconid Opius phaseoli, 

is known from eastern Africa, India and the 

Philippines and has been introduced to Hawaii and Taiwan. This species is capable of 

parasitisation levels of up to 90% or more and, when introduced to Hawaii, it and the related 

O. importatus resulted in successful biological control of bean fly. 

There are good reasons for countries where bean fly is a problem and where 

parasitisation levels are low, to consider introducing these and other parasitoids to assist in 

reducing bean fly populations. 

20° 

0° 

20°


Ophiomyia phaseoli (Tryon) 

Rating 

Origin 

Distribution 

Diptera: Agromyzidae (this species was earlier included in the 

genus Agromyza or Melanagromyza) 

bean fly 


+++ Indo +++ Guam 

14 ++ Laos, Viet, Msia, 

Phil 

9 ++ Fiji, PNG 

+ Myan, Thai, Sing + Sam, Sol Is 

P Brun P P FSM 

Unknown, but presumably in association with one of its current legume host 

genera in India or possibly Southeast Asia. It was described by Tryon (1895) 

from specimens causing damage to beans in 1888 in Queensland, Australia 

and, shortly after, reported to cause similar damage in New South Wales 

(Froggatt 1899). 

This was given (CIE 1974a) as Africa: 

Burundi, Congo (Zaire), Egypt, 

Ethiopia, Kenya, Madagascar, Mauritius, Malawi, Mali, Nigeria, RŽunion, 

Rwanda, Senegal, South Africa, Sudan, Tanzania, Uganda, Zambia, 

Zimbabwe; Asia: 

Bangladesh, China, Hong Kong, India, Indonesia, Iraq, 

Israel, Jordan, Malaysia, Myanmar, Nepal, Pakistan, Philippines, Ryukyu Is, 

Singapore, Sri Lanka, Taiwan, Thailand; Australia and Pacific Islands: 

Australia, Caroline Is, Fiji, Hawaii, Irian Jaya, Mariana Is, Papua New 

Guinea and Samoa. To the above must be added Israel (Spencer 1990) 

Brunei, Laos, Singapore (Waterhouse 1993b) and Japan (Makino et al. 

1990). It has not been recorded from Europe or the Americas. 

A morphologically very similar species, O. spencerella, 

only readily 

distinguishable from O. phaseoli by the male genitalia, occurs in association 

with it in Kenya, Uganda, Tanzania and Nigeria on Phaseolus vulgaris and, 

less commonly, on several other legumes. There are 3 economically 

important agromyzid miners other than Ophiomyia phaseoli that attack 

much the same legumes in Asia. Melanagromyza (= Agromyza) 

obtusa is 

widely distributed in India as a pest of the developing seeds of chick and 

pigeon peas. The stem miners M. sojae and M. dolichostigma are pests of 

soybean in Indonesia, Japan and Taiwan and damage French beans and 

cowpeas in Sri Lanka and East Africa (Singh and van Emden 1979).

Biology 

Host plants 

4.13 


237 

The adult O. phaseoli is a small fly (females 2.2 mm and males 1.9 mm in 

length), shiny black in colour except for legs, antennae and wing veins, 

which are light brown (Abul-Nasser and Assem 1966). Females generally 

oviposit in bright sunlight in the upper surface of the cotyledons (soybeans) 

or young leaves of its many hosts, laying from 100 to 300 eggs during a 

2Ðweek period (Otanes y Quesales 1918). Not all ovipositor punctures 

receive an egg, many provide sap which the females ingest (Goot 1930). On 

hatching from the egg after 2 to 4 days, the young larva forms a short leaf 

mine before tunneling into the nearest vein. Next the petiole is mined and the 

larva then moves down the stem (Taylor 1958). In young plants the main 

feeding takes place in the lower layers of the stem and the tap root may be 

penetrated. When larvae are numerous, some feed more deeply inside the 

stem and higher up in the plant. 

The larval and pupal stages occupy 7 to 10 days and 9 to 10 days 

respectively, resulting in a life cycle of about 3 weeks (Taylor 1958; Ooi 

1988). However, the life cycle may be as short as 17 days in the field in 

Malaysia (Khoo et al. 1991) and in the laboratory in India at 24¡ to 31¡C as 

short as 11 days (Singh et al. 1991). At the other end of the scale, at higher 

altitudes in Java, the larval stage can be extended from 17 to 22 days and the 

pupal stage from 13 to 20 days (Goot 1930). Pupation occurs head upwards 

beneath the epidermis and generally near the base of the stem (Greathead 

1969). In older plants, larvae may pupate at the base of the petioles. 

Talekar and Lee (1989) have developed a method for mass rearing bean 

fly on newly-germinated soybean cotyledons, permitting one person to 

produce 2 000 adults per day. 

Bean fly is known to attack at least 40 plant species. Most of its important 

hosts belong to the legume tribe Phaseoleae and particularly to the genus 

Phaseolus. 

The very susceptible P. vulgaris (French, kidney, haricot, runner 

or snap bean) is of Central American origin as are several other economic 

species of Phaseolus. 

However, from the point of view of the possible origin 

of O. phaseoli in Asia, all of the Asiatic species formerly placed in the genus 

Phaseolus have now been placed in the genus Vigna (Verdcourt 1970) and it 

is relevant that a number of important Vigna species are believed to have 

originated in India or nearby (Purseglove 1968). These include 

V. aconitifolia (moth bean), V. aurea (green or golden gram, mung bean) 

V. calcarata (rice bean) and V. mungo (black gram, urd bean). Other 

important hosts include Cajanus cajan (pigeon pea: origin Africa); Glycine


Damage 

max (soybean: southern China); Lablab niger (= Dolichos lablab) 

(hyacinth 

bean: India); Pisum sativum (pea: southwestern Asia); and Vigna 

unguiculata (cowpea: Africa) (Purseglove 1968; Spencer 1973). Wild hosts 

include Canavalia ensiformis, 

Crotalaria juncea, 

C. laburnifolia, 

C. mucronata, 

Macroptilium atropurpureum, 

M. lathryoides, 

Phaseolus 

panduratus, 

P. semierectus and Vigna radiata (Goot 1930; Kleinschmidt 

1970; Spencer 1973; Abate 1991). 

There is a large variation in susceptibility between different cultivars of 

susceptible species. This variation affords an important opportunity to select 

cultivars that suffer comparatively little damage and is being extensively 

investigated (e.g. Annappan et al. 1984, Gill and Singh 1988; AVRDC 1990, 

Talekar and Hu 1993; Talekar and Tengkano 1993; Gupta et al. 1995). Both 

morphological and chemical characteristics are involved (Chiang and Norris 

1983). 

O. phaseoli can be a limiting factor in the cultivation of susceptible legumes 

in Southeast Asia and most other regions where it occurs. Spencer (1973) 

and many other authors consider it to be one of the most serious of all 

agromyzid pests. Losses of 50% to 100% of crops are reported from many 

parts of the world, and are particularly heavy under dry conditions. 

Although some leaves may wilt as a result of mining and petiole 

tunnelling, most damage results from destruction of tissue at the junction of 

the stem and root. When damage is limited, plants may survive by forming 

adventitious roots (e.g. with P. vulgaris and soybeans) and produce a limited 

crop. However, seedlings most frequently die. Plants that do not respond 

rapidly to root damage and develop adventitious roots are liable to break off 

at ground level during windy periods. When infestations are heavy, the 

aggregation of puparia within the stem results in it swelling, splitting open 

and rotting. De Meijere (1922) reported that young plants in Sumatra 

generally died when they contained 10 to 20 larvae. In Egypt, 25 larvae and 

pupae have been found in a single bean plant (Hassan 1947) and as many as 

320 in a cowpea plant (Abul-Nasr and Assem 1966). 

Seed treatments or post emergence sprays with broad spectrum 

insecticides have been used to control O. phaseoli but, inter alia, 

they 

undoubtedly have serious adverse effects on its parasitoids. Useful control 

has been obtained by intercropping, the use of resistant varieties, adjusting 

planting dates, crop rotation and other cultural methods such as planting into 

rice stubble or covering the newly sown areas with rice straw. It is clear, 

however, that, in areas where effective parasitoids are already present, these 

can play an important role in minimising bean fly damage if not interfered 

with by insecticides.


4.13 


More than 50 parasitic Hymenoptera have been reported from bean fly 

(Table 4.13.1), almost all emerging from pupae, arising from eggs laid in 

host larvae. No egg parasitoids are known and no dipterous parasitoids have 

been recorded. Although figures for percent parasitisation are unavailable 

from many countries where bean fly occurs, the levels recorded are 

generally unimpressive, often less than 30%. The outstanding exception is 

the 90% or more, often produced by the braconid Opius phaseoli in East 

Africa and Ethiopia (Greathead 1969; Abate 1991) and similar levels from 

Opius phaseoli and Eurytoma sp. in the Agra region of India (Singh 1982). 

AUSTRALIA 

Although Tryon (1895), who described Ophiomyia phaseoli from 

Queensland specimens, believed it to be native to Australia, it has never been 

recorded extensively from indigenous plants (Kleinschmidt 1970), so is 

unlikely to have evolved there. Twelve parasitoids (two of them possibly 

hyperparasitoids) (Table 4.13.1) emerged from O. phaseoli pupae taken 

mainly from cowpea ( Vigna unguiculata). 

A later re-examination of 

Australasian Chalcidoidea by Bou‹ek (1988) indicates that only 11 species 

were actually involved and none were hyperparasitoids. Bou‹ek (1988) lists 

one additional parasitoid, the eurytomid Plutarchia bicarinativentris, which 

also occurs in Papua New Guinea. The braconid Opius oleracei, which 

attacks bean fly, has also been recorded from the widespread agromyzid 

Chromatomyia horticola (= Phytomyza atricornis). 

EAST AFRICA 

Ophiomyia phaseoli was not reported in Uganda until the 1920s, Tanzania 

until 1937 and Kenya until 1939. It occurs in close association with 

Ophiomyia spencerella and also with O. centrosematis, 

a species which is 

known also from Indonesia. 

The bean fly is very heavily attacked in all climatic zones by the braconid 

Opius phaseoli (usually of the order of 70% to 90% and even up to 94.4% 

parasitisation), and relatively lightly (3% to 9%) by Opius importatus, 

by a 

polyphagous pteromalid near Herbertia sp. (less than 1%) and by an 

assemblage of chalcidoids. Puparia are also consumed by ants. Three wasps 

found in small numbers were considered to be hyperparasitoids, namely 

Norbanus sp. (Pteromalidae), Eupelmus sp. nr australiensis (= E. sp. nr 

popa) 

(Eupelmidae) and Pediobius sp. (Eulophidae). In spite of the heavy 

parasitisation by Opius phaseoli, 

sufficient bean flies survive in some 

seasons to cause heavy infestations of plantings. These parasitoids are thus, 

at times, unable to prevent economic damage (Greathead 1969). 

239

Table 4.13.1 

Natural enemies of Ophiomyia phaseoli 

Species 

HYMENOPTERA 

BRACONIDAE 


Fopius sp. Thailand Burikam 1978; Burikam & Napompeth 1979, 

Napompeth 1994 

Opius importatus 

East Africa 

Hawaii 

Opius ?liogaster* Mauritius 

Zimbabwe 

Greathead 1969, 1975 

Raros 1975 

Moutia 1932 

Jack 1913; Taylor 1958 

Opius oleracei Australia Kleinschmidt 1970 

Opius phaseoli Botswana 

East Africa 

Ethiopia 

Hawaii 

India 

Madagascar 

Malawi 

Mauritius 

Philippines 

Taiwan 

Zimbabwe 

Opius sp. 

CHALCIDIDAE 

Taiwan Chu & Chou 1965 

1. East Africa Greathead 1969 

2. Sri Lanka Rutherford 1914b 

3. 

* probably Opius phaseoli (Fischer 1971a) 

Zimbabwe Jack 1913 



Abate 1991; Greathead 1969 

Davis 1971, 1972; Fischer 1971a; Greathead 1975; 

Raros 1975 

Fischer 1963; Ipe 1987 


Letourneau 1994 


Fischer 1971b 

N.S. Talekar pers. comm. 1994 



Table 4.13.1 (contÕd) Natural enemies of Ophiomyia phaseoli 

Species 

HYMENOPTERA 


CYNIPIDAE 

Cynipoide sp. Indonesia Goot 1930 

Eucoilidea sp. Taiwan Chu & Chou 1965 

Unidentified Thailand Burikam 1978 

EULOPHIDAE 

Aprostocetus sp. Ethiopia Abate 1991 

Chrysonotomyia douglasi Australia Bou‹ek 1988; Kleinschmidt 1970 

Chrysonotomyia sp. nr erythraea Ethiopia Abate 1991 

Chrysonotomyia formosa Ethiopia Abate 1991 

Chrysonotomyia sp. Australia Dodd 1917 

Cirrospilus sp. Ethiopia Abate 1991 

Euderus sp. Thailand Burikam 1978; Napompeth 1994 

Hemiptarsenus varicornis Australia Dodd 1917; Kleinschmidt 1970; Bou‹ek 1988 

Hemiptarsenus sp. Australia 

Philippines 

Kleinschmidt 1970 

Litsinger 1987 

Meruana liriomyzae Ethiopia Abate 1991 

Pediobius acantha Ethiopia Abate 1991 

Pediobius sp. East Africa Greathead 1969 

Tetrastichus sp. India Gangrade 1974; Ipe 1987 

Unidentified Hawaii Raros 1975 

4.13 Ophiomyia phaseoli 241


Species 

HYMENOPTERA 


EUPELMIDAE 

Eupelmus ?australiensis Ethiopia, East Africa Greathead 1969; Abate 1991 

Eupelmus grayi Australia Kleinschmidt 1990 

Eupelmus sp. nr urozonus Egypt 

Ethiopia 

Eupelmus sp. Australia 

Ethiopia 

EURYTOMIDAE 

Eurytoma larvicola Australia 

Egypt 

Eurytoma poloni Indonesia 

Malaysia 

Philippines 

Eurytoma spp. 2 ´ 

1 ´ 

1 ´ 

1 ´ 

1 ´ 

1 ´ 

Australia 

Egypt 

Ethiopia 

India 

Indonesia 

Taiwan 

Plutarchia sp. Malaysia 

Philippines 

Abul-Nasr & Assem 1968 

Abate 1991 

Dodd 1917 

Abate 1991 

Kleinschmidt 1970; Bou‹ek 1988 

Hassan 1947 

Goot 1930 

Ho 1967; Yunus & Ho 1980 

Otanes y Quesales 1918 

Dodd 1917; Kleinschmidt 1970 


Abate 1991 

Singh 1982; Ipe 1987 

Goot 1930 

Chu & Chou 1965 

Ooi 1973 


Plutarchia bicarinativentris Australia, PNG Dodd 1917; Bou‹ek 1988 

Plutarchia indefensa Thailand Burikam 1978 



Species 

HYMENOPTERA 


PTEROMALIDAE 

Callitula filicornis Ethiopia Abate 1991 

Callitula viridicoxa Australia, PNG Bou‹ek 1988; Kleinschmidt 1970 

Callitula yasudi Japan Yasuda 1982 

Chlorocytus sp. India Kundu 1985 

Cryptoprymna sp. Egypt 

Taiwan 



Halticoptera ?circulus Ethiopia Abate 1991 

Halticoptera patellana Hawaii Greathead 1975; Raros 1975 

Halticoptera sp. Egypt 

Taiwan 

Herbertia sp. Ethiopia 

East Africa 



Abate 1991 


Oxyharma subaenea Australia Dodd 1917; Kleinschmidt 1970 

Polycystus propinquus Sri Lanka Waterston 1915 

Polycystus sp. India Babu 1977 

Sphegigaster brunneicornis Ethiopia 

India 

Sphegigaster hamygurivara Japan Yasuda 1982 

Sphegigaster rugosa India 

Sri Lanka 

Abate 1991 

Peter & Balasubramanian 1984 

Ipe 1987 

Waterston 1915 

4.13 Ophiomyia phaseoli 243


Species 

HYMENOPTERA 


PTEROMALIDAE (contÕd) 

Sphegigaster stella Malaysia 

Philippines 

Sphegigaster stepicola Ethiopia Abate 1991 

Sphegigaster voltairei Australia, PNG 

Egypt 

Indonesia 

Sphegigaster sp. Japan 

Philippines 

Taiwan 

Ho 1967 

Otanes y Quesales 1918 

Syntomopus shakespearei Australia Kleinschmidt 1970 

Syntomopus sp. Japan Yasuda 1982 

Unidentified Thailand Burikam 1978 

TETRACAMPIDAE 

Epiclerus sp. nr nomocerus Ethiopia Abate 1991 

A NEMATODE East Africa 

Thailand 

Dodd 1917; Kleinschmidt 1970; Bou‹ek 1988 

Hassan 1947 

Goot 1930 

Yasuda 1982 




Burikam 1978 


EGYPT 

ETHIOPIA 

4.13 Ophiomyia phaseoli 245 

The bean fly was first reported in 1922, together with a parasitoid, later 

identified as Sphegigaster voltairei (= Trigonogastra agromyzae) 

(Pteromalidae). The eurytomid Eurytoma larvicola was also recorded 

(Hassan 1947). Later, Abul-Nasr and Assem (1968) recorded 5 parasitoid 

species emerging from bean fly puparia in the laboratory (Table 4.13.1), but 

no information was provided on their effectiveness. 

O. phaseoli was first reported in the early 1970s from Phaseolus vulgaris 

(French bean), Vigna unguiculata (cowpea) and soybean (Glycine max), 

although economic damage occurred only on French bean. The leguminous 

bush, Crotalaria laburnifolia was the only wild host which supported bean 

fly and its 17 species of parasitoids throughout the year (Table 4.13.1). Of 

these parasitoids, the pteromalids Sphegigaster stepicola and 

S. brunneicornis were the commonest species, accounting for up to 44.5% 

(average 26.2%) of total parasitisation on Crotalaria. Parasitisation by the 

braconid Opius phaseoli averaged a low 5.6% (range 0% to 23.2%) on 

Crotalaria, but it was the major parasitoid on French bean, accounting for 

over 87% of the total parasitisation. Abate (1991) concluded from this that 

the host plant plays an important role in Ophiomyia phaseoli population 

dynamics, although it remains to be shown whether this applies equally to 

the range of food legumes. It is, perhaps, relevant that the bean fly acts as a 

true leaf miner on Crotalaria, the larvae mining and eventually pupating 

within the leaf. This behaviour renders both stages more accessible to 

smaller parasitoids than when larvae are in the deeper tissues of a bean stem. 

The remaining 14 parasitoids recorded from Ethiopia (Table 4.13.1) were 

classified as very rare, (defined by Abate (1991) as equal to or less than 10% 

of total insect emergence) and together on average caused 9.6% mortality. 

Observations suggested that mortality of French bean seedlings caused by 

bean fly was much less severe in areas where the wild host, Crotalaria, 

occurred (14.8% in 1987 and 3.8% in 1988) than in its absence (39.1% in 

1987 and 36.1% in 1988 (Abate 1991)). This is circumstantial evidence that 

natural enemies play an important part in regulating bean fly populations. 

Negasi and Abate (1986) recorded a pteromalid Cyrtogaster sp. from 

O. phaseoli on French beans but, as this was not mentioned in the later, much 

more detailed paper on parasitoids by Abate (1991), it has not been included 

in Table 4.13.1.


HAWAII 

Damaging populations of O. phaseoli built up rapidly and caused 

widespread damage to cultivated legumes after it was first recorded in 1968. 

The only parasitoid found attacking it at that stage was the pteromalid 

Halticoptera patellana, a polyphagous European parasite of agromyzids 

(Greathead 1975). 

INDIA 

Ophiomyia phaseoli is said not to cause as serious damage to food legumes 

in India as it does elsewhere, especially in Indonesia and East Africa 

(Talekar 1990). 

The pteromalid Chlorocytus sp. was reported to parasitise 8% to 10% of 

O. phaseoli puparia infesting stems of soybean (Glycine max) in the New 

Delhi area (Kundu 1985). In Bangalore, the pteromalid Sphegigaster 

brunneicornis, which emerged from bean fly puparia in cowpea, was the 

only parasitoid recorded. The extent of parasitisation ranged from 16.7% in 

July to 85.5% in September, but this did not achieve adequate control 

because bean fly infestation rose from 12% in July to 68% in September 

(Peter and Balasubramanian 1984). Ipe (1987) recorded four parasitoids (a 

eurytomid, Opius phaseoli, Tetrastichus sp. and Sphegigaster rugosa) from 

Agra. He commented that Ophiomyia phaseoli infestations are kept under 

control by parasitoids and that the percentage of parasitisation reaches 

appreciable levels each season. Babu (1977) reported Polycystus sp. 

emerging from bean fly puparia and causing parasitisation ranging from 

3.4% during February to 61% in August. However, the highest levels of 

attack are those recorded at Agra by Singh (1982) for Opius phaseoli and 

Eurytoma sp.. The parasitisation of bean fly infesting cowpea reached 46.2% 

during early October, rising to 94% by the end of November. From 

December to March these parasites effectively controlled Ophiomyia 

phaseoli populations infesting cowpea, garden pea and Lablab niger (Singh 

1982). 

It appears that the different species of bean fly parasitoids in India may 

not occur at all widely. Whether this is due to climatic limitations, sampling 

from different hosts, or overall inadequate sampling of populations remains 

to be determined. 

INDONESIA 

The most comprehensive account of the biology, hosts and parasitoids of 

bean fly in Indonesia was published in Dutch by Goot (1930). This was 

translated into English and republished in 1984 by the Asian Vegetable 

Research and Development Center Taiwan. A summary appears in 

Kalshoven (1981).


Three agromyzids are pests of soybean and some other economically 

important legumes in Indonesia: Ophiomyia phaseoli (by far the most 

important), the soybean stem borer Melanagromyza sojae (which bores into 

the pith of the stem, but seldom kills the plant) and the soybean top borer 

M. dolichostigma (which bores into the tops and causes stunting). The first 

two species have been known in Java since 1900. 

Four bean fly parasitoids are known (Goot 1930: Table 4.13.1), but their 

combined effect is generally low, parasitisation averaging 5.1%, with a 

maximum of 42.4%. All species emerge from the host pupa. The most 

effective species is the pteromalid Sphegigaster voltairei, which comprised 

59.1% of the parasitoids. It is also the most important parasitoid of 

Melanagromyza sojae and occasionally attacks M. dolichostigma. Next in 

importance is the cynipid Cynipoide sp., contributing 10.5%: it also attacks 

the two other agromyzids. Finally, Eurytoma poloni and Eurytoma sp. each 

contribute 0.2% to the total. Both are more frequently bred from the two 

other agromyzids. 

MALAYSIA 

The bean fly was first reported in Peninsular Malaysia in 1924 and is 

regarded as the most important pest of green gram (Phaseolus aureus). It can 

also cause serious damage to French bean and other legume crops. Two 

parasitoids, which also occur in the Philippines, are Eurytoma poloni 

(Eurytomidae) and Sphegigaster (= Paratrigonogastra) stella (Ho 1967). 

Ooi (1973) recorded the eurytomid Plutarchia sp. 

PHILIPPINES 

The bean fly was first noticed in 1912, but was thought at the time to have 

been present for some years. Two parasitoids are known, a more abundant 

Eurytoma poloni (Eurytomidae) and a less abundant Sphegigaster stella, in 

the ratio 60:47. Their joint parasitisation averaged 17% with a range from 

6% to 49% (Otanes y Quesales 1918). Opius phaseoli was not recorded in 

this study. 

SRI LANKA 

Ophiomyia phaseoli was first recorded in 1901. Several unidentified species 

of hymenopterous parasitoids were bred by Rutherford (1914b) but, 

although they Ôno doubt do a considerable amount of goodÕ, they were 

unable to keep the fly in check. 

TAIWAN 

Chu and Chou (1965) reported a braconid (Opius sp.), 4 pteromalids 

(Cryptoprymna sp., Halticoptera sp., Sphegigaster sp. and Eucoilidea sp.) 

and a eurytomid (Eurytoma sp.) parasitoid attacking bean fly infesting 

soybean and Rose et al. (1976) added another eurytomid (Plutarchia sp.).


Chiang et al. (1978) surveyed the parasitoids of three agromyzids 

infesting mungbean, namely Ophiomyia phaseoli, O. centrosematis and 

Melanagromyza sojae. Parasitisation fluctuated considerably, surpassing 

60% during July, but declining to nearly 0% in December and January. Since 

there was a negative correlation between agromyzid populations and percent 

parasitisation it was concluded that parasitoids played a role in controlling 

agromyzid populations. 

THAILAND 

Surveys in several regions of Thailand for natural enemies of the major pest 

of soybean (Ophiomyia phaseoli) revealed 5 species of hymenopterous 

parasitoid and a nematode (Burikam 1978). The most important species 

were Plutarchia indefensa (Eurytomidae) and Fopius sp. (Braconidae). A 

eulophid, a pteromalid and a cynipid were less important (Burikam and 

Napompeth 1979; Napompeth 1994). Parasitisation by P. indefensa 

averaged 52.8% and two samples gave 7.5% and 5.9% for Fopius sp. 

(Burikam 1978), although Napompeth (1994) later considered the two 

species to be of equal importance. The cynipid wasp also attacked the bean 

stem miner Melanagromyza sojae. On one occasion pupae containing 20 to 

50 nematodes were recorded, with a pupal parasitisation rate of 4.6% 

(Burikam 1978). A life table analysis showed that there was a densitydependent 

factor regulating bean fly populations (Burikam and Napompeth 

1979). 

Unless Fopius sp. (Braconidae) proves to be Opius phaseoli 

(Braconidae) (which is present in both India and the Philippines), it appears 

that O. phaseoli may not be widespread throughout Southeast Asia and may 

be well worth distributing more widely. The identity of the Fopius sp. is to 

be investigated (B. Napompeth pers. comm. 1994). 

ZIMBABWE 

Phaseolus spp. are the principal host crops damaged, but cowpeas (Vigna 

sinensis) and soybeans (Glycine soja) are also attacked. Plantings in late 

summer are usually only lightly infested and a braconid, identified as Opius 

liogaster (but quite possibly O. phaseoli), exercises effective control in most 

years (Taylor 1958). Earlier Jack (1913) had reported that a braconid larval 

parasitoid was ineffective in controlling bean fly although it was bred freely 

from O. phaseoli late in the season.



The only introductions for biological control of bean fly (Table 4.13.2) have 

been of the two braconid parasitoids Opius phaseoli and O. importatus from 

Uganda to Hawaii in 1969 (Davis 1971, 1972; Greathead 1975; Funasaki et 

al. 1988) and of O. phaseoli from Hawaii to Taiwan in 1974Ð75 (N.S. 

Talekar pers comm. 1994). In Hawaii both species rapidly became 

established on Oahu and host density was soon markedly reduced. They 

were introduced to other islands and, by 1971, on Kauai 100% of bean flies 

sampled produced parasitoids: on Maui rates ranged from 25% to 83%. No 

differences in the incidence of parasitisation were detected when 

infestations on French bean and cowpea were compared. By 1973 

O. importatus had become the dominant parasitoid and the polyphagous 

Halticoptera patellana (which was already present) was only rarely 

encountered (Greathead 1975). Surveys by Raros (1975) in 1973 and 1974 

of three locations on Oahu revealed average parasitisation ranging from 

8.3% to 23.5%, a result Talekar (1990) suggested might have been due to the 

heavy use of insecticides diminishing the earlier effectiveness of the 

parasitoids. In 1994 the bean fly was reported to be still a problem on young 

seedlings, so that farmers usually apply one or two insecticide sprays after 

seedlings emerge above the ground. Once the bean plant has developed a 

couple of leaves the fly is no longer a problem (W.C. Mitchell pers. comm.). 

Opius phaseoli was not recovered in Taiwan for the first two years after its 

introduction in 1974Ð75 from Hawaii, but there have been recent reports of 

its presence, in spite of the current excessive use of insecticides against bean 

fly (N.S. Talekar pers. comm. 1994).

Table 4.13.2 Introductions for the biological control of Ophiomyia phaseoli 

Species 

BRACONIDAE 

Origin Liberated Year Result Reference 

Opius importatus Uganda Hawaii 1969 + Fischer 1971a; Davis 1971, 1972; Funasaki et 

al. 1988; Greathead 1975; Raros 1975 

Opius phaseoli Uganda 

Hawaii 

Hawaii 

Taiwan 

1969 

1974Ð75 

+ 

+ 

Fischer 1971a; Raros 1975 

N.S. Talekar pers. comm. 1994 


The more important parasitoids 


The names of a number of species are now different from those used by 

earlier authors. To enable cross referencing with those used in Table 4.13.1, 

the older names are shown below, together with a summary of information 

available on the biology of the more important species. 

Callitula viridicoxa (= Eurydinotellus viridicoxa = Polycystomyia 

beneficia) Hym.: Eurytomidae 

Chrysonotomyia (= Achrysocharis) douglasi Hym.: Eulophidae 

Chrysonotomyia ?erythraea Hym.: Eulophidae 

Chrysonotomyia formosa Hym.: Eulophidae 

The two latter species are widely distributed primary parasitoids attacking 

bean fly infesting Crotalaria in Ethiopia, parasitisation ranging from 0% to 

8.7% (average 2.6%). C. formosa has also been recorded from Liriomyza 

trifolii infesting beans in Guam (Schreiner et al. 1986; Abate 1991). 

Cynipoide sp. Hym.: Cynipoidea 

This parasitoid has only been reported from Java, where Goot (1930) found, 

from 90 samplings between 1919 and 1923, that it constituted 40% of the 

parasitoids reared, although the level of parasitisation varied greatly. It also 

emerged from puparia of Melanagromyza sojae and M. dolichostigma. It 

occurred in both tropical lowland and cool highland conditions. 

Euderus sp. Hym.: Eulophidae 

A tentative assignation as Euderus ?sp. was made by Napompeth (1994) of 

the tiny eulophid recorded by Burikam (1978) and Burikam and Napompeth 

(1979). More than one parasitoid could be produced per bean fly host. The 

female parasitoid oviposited in the first instar host larva and pupation 

occurred during the hostÕs third instar, either within or alongside the host. 

The pupal stage averaged 7 days (Burikam 1978). 

Eurytoma poloni Hym.: Eurytomidae 

This parasitoid has been recorded from Indonesia, Malaysia and the 

Philippines, but little is known of its biology. It was recorded only once in 

Java from Ophiomyia phaseoli so it is clearly not an important parasitoid of 

the bean fly there. In fact, it generally emerges from Melanagromyza 

dolichostigma. Adults live 22 to 28 days (Goot 1930). 

Fopius sp. Hym.: Braconidae 

This was referred to as Biosteres sp. by Burikam (1978) and Burikam and 

Napompeth (1979), but altered to Fopius sp. by Napompeth (1994). Adults 

emerged from bean fly puparia (one per host) and mated on the first day. 

Within two days 27 mature and immature eggs could be counted per female.


Two samples in August revealed parasitisation levels of 5.9% and 7.5% 

(Burikam 1978). 

Halticoptera ?circulus Hym.: Pteromalidae 

This is a widespread primary parasitoid of agromyzid leafminers in many 

parts of the world. It was recorded as very rare (² 10% parasitisation) on 

bean fly in Ethiopia (Abate 1991). 

Hemiptarsenus varicornis (= Neodimmockia agromyzae and 

probably = Hemiptarsenus semialbicornis) Hym.: Eulophidae 

This is a very widespread parasitoid of dipterous leaf miners, including 

O. phaseoli and Liriomyza sativae. It occurs throughout tropical and 

southern temperate countries of the eastern hemisphere. In Australia it is 

common in many places along the eastern and southeastern coast. It also 

occurs in New Zealand, New Caledonia, Fiji and Vanuatu; Malaysia, Sri 

Lanka, India, Pakistan and Saudi Arabia; Senegal, Ghana, Sudan, Ethiopia, 

Kenya and Tanzania (Bou‹ek 1988). 

Meruana liriomyzae Hym.: Eulophidae 

This species was recorded as rare (³10% parasitisation) on bean fly infesting 

Crotalaria in Ethiopia (Abate 1991). It is also known from Liriomyza 

brassicae in Mauritius, L. sativae in Mauritius and RŽunion, from L. trifolii 

in Kenya, Chromatomyia horticola (= Phytomyza atricornis) in Ethiopia 

and South Africa, and from unidentified hosts in Australia and Zimbabwe 

(Bou‹ek 1988; Abate 1991). 

Opius importatus Hym.: Braconidae 

This species was first recorded from East Africa as Opius sp. by Greathead 

(1975) and later described as O. importatus by Fischer (1971b). In nature, it 

is known only from East Africa and only from Ophiomyia phaseoli. When 

first taken between November 1967 and April 1968 it attained parasitisation 

levels of 3% to 9% (Greathead 1969). However, later samples taken in 

Uganda in 1971 contained nearly 50% of O. importatus (Greathead 1975). 

The first instar larva was found in the third instar host larva and developed 

rapidly once the host pupated. Adults, that are similar in appearance to dark 

specimens of Opius phaseoli, emerge about 33 days after the appearance of 

the host plant above the soil. O. importatus was inadvertently included in 

shipments of Opius phaseoli to Hawaii, where it soon became the dominant 

parasitoid of Ophiomyia phaseoli (Greathead 1975). 

Opius phaseoli (= O. melanagromyzae) Hym.: Braconidae 

O. phaseoli was originally described from the Philippines (Manila) without 

a host by Ashmead (1904) as Eurytenes nanus. However, as this name was 

preoccupied, it was redescribed as O. phaseoli by Fischer (1963), who listed


its host as Ophiomyia phaseoli and its distribution as India (Nagpur) and the 

Philippines. Later, Fischer (1966) listed it as a parasitoid of the leaf miner 

Melanagromyza atomella in India and Singh (1982) from Melanagromyza 

sojae. Both Singh (1982) and Ipe (1987), working in the area of Agra, 

considered it to be important in regulating bean fly populations there. No 

information is available on its presence or effectiveness elsewhere in Asia or 

Southeast Asia, except for the original record of a single female wasp from 

Manila (Ashmead 1904). 

In East Africa, Greathead (1969, 1975) reported parasitisation levels of 

Ophiomyia phaseoli by Opius phaseoli that were frequently above 50%, and 

sometimes reached 94.4%. Levels of up to 10% on Ophiomyia spencerella 

were also recorded and Opius phaseoli was twice reared from Ophiomyia 

centrosematis. He concluded that Opius phaseoli was the chief biotic factor 

limiting the population of bean fly in East Africa. Nevertheless, he reported 

levels of only 38% parasitisation from 3 bean fly samples taken in 

Madagascar and only about 20% in Mauritius. He suggested that the latter 

result might be due to a different strain of the parasitoid or, perhaps, even a 

different species. In Ethiopia Abate (1991) reported over 93% parasitisation 

of bean fly on French bean, but much lower levels (average 5.6%) on a wild 

host Crotalaria. 

One to five (but generally two or three) eggs are laid at a time, usually in 

first instar host larvae. Hatching occurs about 2 days later, soon after the host 

larva has moulted to the second instar. The first instar parasitoid larva grows, 

but does not moult until its host has pupated. Meanwhile, all except one 

Opius larva are suppressed. Development is rapid in the host pupa, the entire 

larval period lasting 9 to 10 days, leading to a prepupal period of 1 to 2 days 

and a pupal period (within the host puparium) of about 3 days (Raros 1975). 

In East Africa, the pupal period lasts a minimum of 4 days and adults 

commence emerging about 30 days after the appearance of the host plant 

above the ground (Greathead 1969). On average, males live 20 and females 

23 days. Females mate a day after emergence and, following a 

preoviposition period of 1 to 2 days, may lay up to 358 eggs throughout life. 

First instar host larvae are preferred to second instar larvae (62.9:32.8) 

(Raros 1975). Other host stages are not attacked. Adult parasitoids feed on 

water droplets and host plant sap resulting from oviposition punctures made 

either by host adults or by female parasitoids. The male:female sex ratio was 

about 1:1.4 in Hawaii (Raros 1975).


Oxyharma (= Pterosema) subaenea Hym.: Pteromalidae 

Plutarchia indefensa Hym.: Eurytomidae 

This wasp was referred to by Burikam (1978) and Burikam and Napompeth 

(1979) as Plutarchia sp. Eggs are laid in third instar larvae of Ophiomyia 

phaseoli, usually at the posterior end. One or two eggs are laid per host and, 

on hatching after 2 to 3 days, one surviving larva remains in the first instar 

until the host has pupated. The larval stage lasts 5 to 7 days and the pupal 

stage (inside the host puparium) 7 to 8 days, giving a development period 

from egg to adult of 16 to 19 days. Adult males lived 4 to 19 days (average 

11.5) and females 10 to 25 days (average 16.9), during which 6 to 14 eggs 

developed per day (Burikam 1978). 

Pteromalid Hym.: Pteromalidae 

The unidentified pteromalid recorded from Thailand (Burikam 1978; 

Burikam and Napompeth 1979; Napompeth 1994) was found parasitising 

pupae of O. phaseoli. The female laid 1 or 2 eggs in the host puparium. These 

hatched in 2 days and, after 4 days larval development, pupation occurred 

within the host puparium. The pupal stage lasted 7 to 8 days, resulting in a 

life cycle of 12 to 14 days. Female wasps lived more than 2 weeks (Burikam 

1978). 

Sphegigaster brunneicornis Hym.: Pteromalidae 

This species has been reported from Ethiopia (Abate 1991), India and Sri 

Lanka (Peter and Balasubramanian 1984). O. phaseoli is its only recorded 

host. 

Sphegigaster (= Trigonogastra) rugosa Hym.: Pteromalidae 

Sphegigaster (= Paratrigonogastra) stella Hym.: Pteromalidae 

Sphegigaster stepicola Hym.: Pteromalidae 

This species is known from Ophiomyia phaseoli in Ethiopia (Abate 1991) 

and from Phytomyza albiceps in southern Europe and India (Abate 1991). 

Combined parasitisation with S. brunneicornis of bean fly on Crotalaria in 

Ethiopia ranged from 3.1% to 44.4% (average 26.2%), of which S. stepicola 

accounted for nearly 72% (Abate 1991). 

Sphegigaster voltairei (= Sphegigaster agromyzae 

= Trigonogastra agromyzae) Hym.: Pteromalidae 

This species is recorded from Australia, Papua New Guinea, Egypt and 

especially Indonesia where Goot (1930) reported that, on average, it 

comprised 60% of emergences from parasitised puparia and that it could be 

kept alive from 30 to 48 days. It was also the most important parasitoid of 

Melanagromyza sojae and was bred several times from M. dolichostigma. 

Syntomopus (= Merismorella) shakespearei Hym.: Pteromalidae

Comment 


Although O. phaseoli occurs in Africa, it now seems unlikely that it evolved 

there. It was reported in Uganda as recently as the 1920s, in Tanzania in 

1937 and in Kenya in 1939. In East Africa, it occurs in close association with 

the very similar O. spencerella, which is generally the dominant species 

(Greathead 1969). Spencer (1973) postulated that O. spencerella evolved in 

Africa and O. phaseoli in Asia. The latter subsequently arrived in Africa to 

occupy a similar niche in much the same host plants, but isolated 

reproductively from O. spencerella. It is interesting that the major natural 

enemy of Ophiomyia phaseoli in East Africa is the braconid Opius phaseoli, 

which also occurs in India, whereas that of Ophiomyia spencerella is a 

cynipid Eucoilidea sp.. Although Eucoilidea sp. is not restricted to 

Ophiomyia spencerella, this species nevertheless does not attack Ophiomyia 

phaseoli. 

In spite of Asia (and probably India) being nominated as the probable 

centre of origin of Ophiomyia phaseoli, it is interesting that bean fly is even 

more heavily parasitised in East Africa and Ethiopia than anywhere else Ñ 

and by 2 braconid parasitoids, one of which is native to the latter regions. 

These are Opius phaseoli and O. importatus. Opius phaseoli was described 

from a single female from Manila, but the absence of records of it attacking 

Ophiomyia phaseoli there or elsewhere in Southeast Asia raises doubts that 

it is native to Southeast Asia. 

The rapid control of Ophiomyia phaseoli in Hawaii following the 

introduction of Opius phaseoli and O. importatus (the latter soon becoming 

the dominant species) demonstrates that, under favourable circumstances, 

biological control alone can produce valuable results. It seems probable that 

this success would be repeated, at least in other Pacific island nations. 

In the far more complex ecological environment in Southeast Asia 

extrapolation from experience in Hawaii is more risky. It is probable that the 

other components of integrated pest management (varietal resistance, 

cultural methods, rational pesticide use, etc.) will all be required to 

supplement the reduction in bean fly density that can be brought about by 

parasitoids. Nevertheless, any substantial decrease in bean fly populations 

that can be achieved by introducing additional effective parasitoids is likely 

to be a valuable contribution towards reduced crop losses.

Table 14 

Table 14 shows for weeds what Table 4 did for 

invertebrates in relation to the top 10 entries. 

Since no information was available on the 

relative rating for the five weeds nominated by 

Tokelau (shown by an asterisk), each was 

70 


allocated the median value of 5. The ranking 

order is only given for species that attain an 

aggregated value of 10 or more. These are 

arranged in descending order of importance in 

Table 15.


71 

Table 14 

The 

relative importance given to the top 10 weeds of agriculture (72 species) of each country in the southern and western Pacific . 

Name Family CoI Fij FrP FSM Gua Kir Mar NCa Niu PNG ASa WSa SoI Tok Ton Tuv Van W.F. No. * Rating Order 

Acacia farnesiana 

Acacia nilotica 

Achyranthes aspera 

Agave americana 

Ageratum conyzoides 

Alternanthera sessilis 

Amaranthus interruptus 

Amaranthus spinosus 

Amaranthus viridis 

( = A. gracilis) 

Antigonon leptopus 

Argemone mexicana 

( = A. americana) 

Bidens alba, Bidens 

pilosa 

Blechum pyrimidatum 

( = B. brownei) 

Brachiaria mutica 

Brachiaria reptans 

Brachiaria 

subquadripara 

Broussonetia papyrifera 

Canavalia rosea 

Cardiospermum 

halicacabum 

Cassytha filiformis 

Cecropia peltata 

Cenchrus echinatus 

Chamaesyce 

( = Euphorbia) hirta 

Chloris barbata 

Mimosaceae 4 1 7 

Mimosaceae 

Amaranthaceae 

Agavaceae 

Asteraceae 2 1 9 

Amaranthaceae 

Amaranthaceae 

Amaranthaceae 

Amaranthaceae 

Polygonaceae 

Papaveraceae 

2 1 9 

Asteraceae 7 9 3 8 1 9 6 29 6 = 

Acanthaceae 8 1 3 

Poaceae 8 1 3 

Poaceae 

Poaceae 

Urticaceae 4 1 7 

Fabaceae 10 1 1 

Sapindaceae 2 1 9 

Lauraceae 7 2 6 3 18 13 

Euphorbiaceae 5 1 6 

Poaceae 3 * 3 3 21 12 

Euphorbiaceae 10 10 * 3 7 45= 

Poaceae

72 


Table 14 

Chromolaena odorata 

Clerodendrum chinense 

( = C. philippinum) 

Clidemia hirta 

Coccinia grandis 

Commelina 

benghalensis 

Commelina diffusa 

Cordia subcordata 

Crassocephalum 

crepidoides 

Crotolaria pallida 

Crotolaria retusa 

Cuphea carthagenensis 

Cynodon dactylon 

Cyperus rotundus 

Dactyloctenium 

aegyptium 

Desmodium incanum 

Digitaria ciliaris 

Digitaria eriantha 

( = D. decumbens) 

Digitaria insularis 

Digitaria setigera 

Echinochloa colona 

Echinochloa crus-galli 

Eichhornia crassipes 

Eleocharis geniculata 

Elephantopus mollis 

( = E. scaber) 

Eleusine indica 

(cont’d) 

Asteraceae 3 7 2 12 21= 

Verbenaceae 2 3 1 3 27 9 

Melastomataceae 4 5 2 13 20 

Cucurbitaceae 5 1 6 

Commelinaceae 2 1 9 

Commelinaceae 

Boraginaceae 

9 6 2 7 


Fabaceae 

Fabaceae 

Lythraceae 

Poaceae 

Cyperaceae 

Poaceae 

7 6 8 7 1 1 5 4 1 1 1 9 1 13 98 1 

Fabaceae 

Poaceae 

Poaceae 

Eleutheranthera ruderalis Asteraceae 

The 



Poaceae 

Poaceae 

Poaceae 

10 1 1 

Poaceae 6 1 5 

Pontederiaceae 

Cyperaceae 

7 1 2 8 4 26 10 

Asteraceae 9 3 2 10 26= 

Poaceae 8 9 6 10 * 10 7 7 21 11


73 

Table 14 

Emilia sonchifolia 

Eragrostis tenella 

Euphorbia heterophylla 

( = E. geniculata) 

Fimbristylis cymosa 

( = F. atollensis) 

Fimbristylis dichotoma 

Fimbristylis miliacea 

Guettarda speciosa 

Hydrilla verticillata 

Hyptis pectinata 

Imperata conferta 

( = I. cylindrica) 

Indigofera suffruticosa 

Ipomoea macrantha 

Ischaemum spp. 

Jatropha gossypifolia 

Kyllinga brevifolia 

Kyllinga nemoralis 

Kyllinga polyphylla 

Lantana camara 

Leucaena leucocephala 

Ludwigia octovalvis 

( = Jussiaea suffruticosa) 

Asteraceae 

Poaceae 

Euphorbiaceae 8 1 3 

Cyperaceae 7 1 4 

Cyperaceae 

Cyperaceae 

Cyperaceae 4 * 2 12 21= 

Hydrocharitaceae 

Lamiaceae 3 1 8 

Poaceae 10 1 1 

Fabaceae 

Convolvulaceae * 5 2 11 23 = 

Poaceae 10 1 1 

Euphorbiaceae 

Cyperaceae 

Cyperaceae 

Macroptilium lathyroides Fabaceae 

Melaleuca 

quinquenervia 

Merremia peltata 

Miconia calvescens 

Mikania micrantha 

Mimosa invisa 

(cont’d) 

The 



Cyperaceae 3 9 2 10 26 = 

Verbenaceae 8 6 6 7 1 6 3 7 40 4 

Mimosaceae 2 8 10 3 13 19 

Onagraceae 5 2 2 15 15 = 

Myrtaceae 

Convolvulaceae 1 10 2 11 23 = 

Melastomataceae 1 1 10 28 = 

Asteraceae 6 9 4 8 6 2 6 7 8 5 10 49 3 

Mimosaceae 1 1 3 4 7 4 3 7 5 2 10 73 2

74 


Table 14 (cont’d) The relative importance given to the top 10 weeds of agriculture (72 species) of each country in the southern and western Pacific . 

Mimosa pigra 

Mimosa pudica 

Miscanthus floridulus 

Momordica charantia 

Monochoria hastata 

Nephrolepis hirsutula 

Ocimum gratissimum 

Oxalis corniculata 

Panicum maximum 

Parthenium 

hysterophorus 


Paspalum conjugatum 

Paspalum dilatatum 

Paspalum paniculatum 

Paspalum vaginatum 

Passiflora foetida 

Passiflora maliformis 

Mimosaceae 3 1 8 

Mimosaceae 5 6 4 6 9 2 6 40 5 

Poaceae 

Cucurbitaceae 

Pontederiaceae 

Davalliaceae 

Lamiaceae 5 1 6 

Oxalidaceae 

Poaceae 3 1 5 


Poaceae 5 8 2 9 

Poaceae 

Poaceae 

Poaceae 5 1 6 

Passifloraceae 

Passifloraceae 

Pennisetum polystachion Poaceae 2 1 9 

Pennisetum purpureum Poaceae 2 1 9 

Phyllanthus amarus Euphorbiaceae 

Physalis angulata Solanaceae 

Pistia stratiotes Araceae 

Pluchea indica Asteraceae 

Portulaca oleracea Portulacaceae 5 3 2 14 18 

Premna obtusifolia 

( = P. serratifolia) 

Verbenaceae 1 1 10 28 = 

Pseudelephantopus 

spicatus 

Asteraceae 

Psidium guajava Myrtaceae 3 1 8 

Ricinus communis Euphorbiaceae 6 1 5

The Major Invertebrate Pests and Weeds of Agriculture and Plantation Forestry in the Southern and Western Pacific 75 


Rottboellia 

cochinchinensis 


Ruellia prostrata Acanthaceae 

Poaceae 5 2 2 15 15 = 

Salvinia molesta Salviniaceae 

Scaveola sericea 

( = S. taccada) 

Goodeniaceae 7 4 2 11 23 = 

Schinus terebinthifolius Anacardiaceae 

Senna ( = Cassia) 

occidentalis 

Caesalpinaceae 

Senna ( = Cassia) tora Caesalpinaceae 8 6 2 8 

Sida acuta Malvaceae 9 9 8 4 4 14 17 

Sida cordifolia Malvaceae 

Sida fallax Malvaceae 8 1 3 

Sida rhombifolia Malvaceae 8 9 7 3 9 

Solanum americanum 

( = S. nigrum) 

Solanaceae 

Solanum mauritianum Solanaceae 

Solanum torvum Solanaceae 2 9 5 1 4 27 8 

Sonchus oleraceus Asteraceae 

Sorghum arundinaceum 

( = S. verticilliflorum) 

Poaceae 4 1 7 

Sorghum halepense Poaceae 9 4 4 3 16 14 

Sorghum sudanense Poaceae 10 1 1 

Spathodea 

companulata 

Bignoniaceae 4 1 7 

Sphaerostaphanos 

invisus 

Thelypteridaceae 

Sphaerostephanos 

unitus 

Thelypteridaceae 

Stachytarpheta 

cayennensis 

Verbenaceae 

Stachytarpheta 

jamaicensis 

Verbenaceae 10 6 2 6

76 D.F. Waterhouse 



Stachytarpheta urticifolia Verbenaceae 10 5 7 10 4 12 20 

Stictocardia tiliifolia Convolvulaceae 

Syndrella nodiflora Asteraceae 

Tecoma stans Bignoniaceae 4 1 7 

Themeda quadrivalis Poaceae 

Tournefortia 

( = Messerschmidia) 

argentea 

Boraginaceae 9 1 2 

Tribulus cistoides Zygophyllaceae 

Tridax procumbens Asteraceae 

Triumfetta rhomboidea Tiliaceae 

Urena lobata Malvaceae 

Vernonia cinerea Asteraceae 9 1 2 

Vigna marina Fabaceae 10 1 1 

Vitex trifolia Verbenaceae 

Wedelia trilobata Asteraceae 

Xanthium pungens Asteraceae 3 1 8

4.15 Planococcus citri 

India 

20° 

Myanmar 

P Laos 

0° 

20° 

China 

++ 

Thailand 

+ 

Cambodia 

Vietnam 

+++ 

+ 

+ Brunei 

Malaysia 

Singapore 

+ 

Indonesia 

Taiwan 

++ 

P 

Philippines 

Australia 

Papua 

New Guinea 

+ 

287 

It is speculated that the citrus mealybug Planococcus citri is of south China origin, 

although it now occurs very widely in tropical, subtropical and temperate regions wherever 

citrus is grown. Like other mealybugs, it is attacked by a large number of non-specific 

predators, especially Coccinellidae, but also Chrysopidae. These consume vast numbers 

of prey when mealybugs are abundant, but often do not reduce host numbers to a level at 

which economic injury no longer occurs. There are several specific or near specific encyrtid 

parasitoids that are capable of lowering P. citri populations below the economic threshold. 

These are worth serious consideration for introduction to regions where they do not already 

occur: Leptomastix dactylopii (of Brazilian origin), Leptomastidea abnormis and Anagyrus 

pseudococci (of Mediterranean origin) and Coccidoxenoides peregrinus (of south China or 

Indian origin). In some situations (particularly cooler conditions) augmentative releases 

are necessary if the use of insecticides is to be avoided. 

20° 

0° 

20°


Planococcus citri (Risso) 

Rating 

Origin 

Distribution 

Hemiptera, Pseudococcidae 

citrus mealybug 

This account draws heavily on that of Bartlett (1978) and CABI abstracts 

since then. 

. 


+++ Viet 

7 ++ ++ 4 

+ Thai, Msia, Brun, 

+ Cook Is, PNG, Tong, 

Indo 

Sam 

P Myan, Phil P Fr P, Niue 

P. citri was described from citrus in southern France (Risso 1813), but 

Bartlett (1978) speculates that it is of Chinese origin. However, the fact that 

it is a widespread and important pest of citrus in 11 of the 14 provinces of 

southern China casts some doubt on this view (Li Li-ying et al. 1997). 

The citrus mealybug is extremely widespread, being present in almost all 

tropical, subtropical and temperate regions of the world and in many 

glasshouses in cooler parts. Other species of Planococcus have often been 

confused with it, including P. pacificus (which does not occur in the 

Mediterranean but is common in the Pacific: Cox 1981, 1989), P. ficus 

(restricted to fig, pomegranate and grape in the Mediterranean: Tranfaglia 

1979; Cox and Freeston 1985; Cox and Ben-Dov 1986) and P. kenyae (on 

Coffea in Kenya: Le Pelley 1943a,b). However, their hymenopterous 

parasitoids discriminate between P. citri and P. kenyae and biological 

control fails when the incorrect parasites are used (Rosen and De Bach 

1977). The collection data of Pacific specimens suggest that P. citri is a 

recent introduction there (Williams 1982). There may well be different 

strains of P. citri. 

For example, in Brazil it is rarely found on citrus, but is 

common on other plants (Compere 1939); and in South Africa it is common 

on citrus, but seldom found on grape vines (De Lotto 1975).

Biology 

Host plants 

4.15 


289 

P. citri shows considerable morphological variation when reared under 

different environmental conditions. Small specimens, produced by rearing 

at high temperatures (32¡C), have smaller appendages and lower numbers of 

cuticular structures than those reared between 17¡ and 25¡C (Cox 1981). 

Adult females are oval, flat and yellow to yellowish brown, with a barely 

visible dorsal line under their waxy covering. Along the edge of the wax 

cover there are short waxy protruberances, the longest of which are at the 

posterior end. Yellow eggs are deposited in an ovisac of wax threads and 

young nymphs are lemon-yellow in colour. Females are oviparous, and 

possibly parthenogenetic when males are not available. 

The development time varies from 20 to 40 days, depending upon the 

host plant and temperature. The pre-oviposition period is 7 to 10 days, eggs 

hatch in 3 to 6 days and, 300 to 500 eggs are laid per female. Additional 

details of developmental periods on coffee and potato sprouts are provided 

by Bartlett and Lloyd (1958) and Martinez and Suris (1987a, b). Eggs, 

nymphs and adults are capable of overwintering. There are 3 instars for 

female nymphs and 2 for male nymphs which then form a waxy puparium. In 

warm areas there are normally 4 or 5 overlapping generations per year 

(Bartlett 1978; Kalshoven 1981), but about double this number under 

laboratory conditions (Gray 1954). 

P. citri can be readily reared in the laboratory on potato sprouts (Fisher 

1963; Martinez and Suris 1987b), lemons and butternut pumpkins 

(Samways and Mapp 1983) and a crawler-proof cage has been described 

(Rao 1989). Dispersal of crawlers is brought about by wind, rain and ants. 

Until mated, females secrete a pheromone continously to attract males: 

(1Rcis) (3-isopropenyl-2-2-dimethylcyclobutyl) methyl acetate (Dunkelblum 

et al. 1986). Synthetic analogues have been tested to reduce 

populations (Rotundo and Tremblay 1974, 1980, 1982; Rotundo et al. 1979). 

P. citri 

has an extremely wide host range, attacking almost every flowering 

plant and some grasses as well. It is most frequently reported from citrus, and 

occurs commonly on other fruit trees and grape vines. Other important crops 

include banana, tobacco, coffee, fig, mango, cocoa, date, casssava, 

macadamia, passionfruit and cut flowers.


Damage 

The citrus mealybug is a widespread and severe pest of citrus; also, in 

temperate areas, of grape vines and, in tropical areas, of coffee and mango. 

P. citri is the most injurious of 6 species of mealybug in the Mediterranean 

basin, the damage caused, especially to fruit, being so severe in France that 

the economic threshold is 2% infestation (Panis 1979). In glasshouses its 

hosts include bulbs, ferns, gardenias and other ornamentals. It is generally 

found on the aerial parts of plants, although a root form occurs. Tender 

growing tips, flower buds and young fruit clusters are favoured aggregating 

points. Excessive removal of sap by large numbers of mealybugs leads to 

wilting and death of shoots and flower buds and to drop of fruit. 

Furthermore, there is abundant growth of sooty moulds on the large amounts 

of honeydew produced, which may render produce unmarketable. P. citri 

becomes most abundant during the dry season. 

P. citri has been incriminated in the transmission of viruses of grape 

vines, cocoa, cucumber, taro and tobacco (Carpenter et al. 1976; Kenten and 

Woods 1976; Bartlett 1978; Legg and Lockwood 1981; Dufour 1988; Agran 

et al. 1990; Pedroso et al. 1991). 


The citrus mealybug, like other mealybugs, is attacked by a wide range of 

naturally occurring predators, particularly coccinellid beetles and lacewings 

(Table 4.15.1). Many of these have a wide to very wide host range and thus 

are less likely nowadays than in the past to be regarded as suitable to 

introduce to new environments. Nevertheless, those reported in recent 

literature are recorded, but no attempt has been made to provide an 

exhaustive list. 

The coccinellid predator Cryplolaemus mountrouzieri in particular has 

been very extensively distributed in the past, often with moderate to good 

success, against a variety of mealybug, scale and aphid pests, including 

P. citri. 

C. montrouzieri and some of the other predators play an important 

role in greatly reducing the abundance of high populations. In general, 

however, their searching ability is poor and prey are missed when population 

densities fall, so their action must generally be supplemented by parasitoids 

or other means for effective control.

Table 4.15.1 

Predators of Planococcus citri 

Species 

HEMIPTERA 

ANTHOCORIDAE 


Orius minutus 

NEUROPTERA 

CHRYSOPIDAE 

Turkey Soylu & Urel 1977 

Brinckochrysa (= Chrysopa) 

scelestes India Krishnamoorthy 1984 

Chrysopa sp. Australia 

USA 

Chrysoperla carnea 

India 

Israel 

USSR 

Mallada (= Anisochrysa) basalis 

Mallada boninensis 

Odontochrysa (= Chrysopa =Plesiochrysa ) 

lacciperda 

Oligochrysa lutea 

Sympherobius barberi 

Sympherobius sanctus 

COLEOPTERA 

COCCINELLIDAE 

Brumoides lineatus 

Brumus suturalis 

Murray 1978 

Meyerdirk et al. 1982 

Krishnamoorthy & Mani 1989b 

Berlinger et al. 1979 

Niyazov 1969 

India Krishnamoorthy & Mani 1989b 

India Krishnamoorthy & Mani 1989b 

India Mani & Krishnamoorthy 1990 

Australia Murray 1978, 1982 

USA Dean et al. 1971; Meyerdirk et al. 1979, 1982 


China Weng & Huang 1988 

Indonesia Kalshoven 1981 

4.15 


291

Table 4.15.1 

Species 

COLEOPTERA 


COCCINELLIDAE (contÕd) 

Chilocorus bipustulatus 

(contÕd) Predators of Planococcus citri 

Israel 

Turkey 

Berlinger et al. 1979 

Soylu & Urel 1977 

Coccinella californica 

California Bartlett & Lloyd 1958 

Coccinella transversalis (= C. repanda) 


Coccinella semipunctata 

USSR Niyazov 1969 

Cryptolaemus affinis 

USA 

Meyerdirk et al. 1979, 1982 

Papua New Guinea Szent-Ivany 1963 

Cryptolaemus montrouzieri 

Easter Is 

Ripa et al. 1995 

India 

Chacko et al. 1978 

Mediterranean 

Panis 1977 

Diomus pumilio 

Australia Meyerdirk 1983 

Exochomus flavipes 

South Africa Samways 1983 

Exochomus flaviventris 

Mediterranean Kanika et al. 1993 

Harmonia octomaculata 

Australia Murray 1978 

Hyperaspis lateralis 

California Bartlett & Lloyd 1958 

Hyperaspis polita 


Hyperaspis 2 ´ spp. USSR Niyazov 1969 

Bennett & Hughes 1959 

Nephus (= Scymnus) bipunctatus 


Nephus (= Scymnus) 

reunioni 

East Africa Ershova & Orlinskii 1982 

Pullus pallidicollis 

India Prakasan 1987 

Scymnus (= Nephus) 

includens 

Italy Tranfaglia & Viggiani 1973 

Scymnus apetzi 


Scymnus apiciflavus 


Scymnus biguttatus 



Table 4.15.1 (contÕd) Predators of Planococcus citri 

Species 

COLEOPTERA 



Scymnus binaevatus South Africa Smith 1923 

Scymnus spp. South Africa 

Turkey 

Samways 1983 


Scymnus roepkei Indonesia Kalshoven 1981 

Scymnus sordidus California Bartlett & Lloyd 1958; 


Scymnus subvillosus USSR Niyazov 1969 

DIPTERA 

CECIDOMYIIDAE 

Coccidodiplosis smithi Indonesia Kalshoven 1981 

Diadiplosis hirticornis Japan Yukawa 1978 

Dicrodiplosis sp. India Chacko et al. 1977 

Triommata coccidivora India Prakasan 1987 

CHAMAEMYIIDAE 

Leucopis alticeps USSR 

Italy 

Niyazov 1969 

Raspi & Bertolini 1993 

Leucopis bella California Bartlett & Lloyd 1958 

Leucopis silesiaca Italy Raspi & Bertolini 1993 

CRYPTOCHETIDAE 

Cryptochetum sp. India Chacko et al. 1977 

SYRPHIDAE 

Syrphus sp. Australia Murray 1982 

4.15 

Planococcus citri 293


Species 

LEPIDOPTERA 


LYCAENIDAE 

Spalgis epius 

HYMENOPTERA 

ENCYRTIDAE 

India Chacko et al. 1977; 

Mani & Krishnamoorthy 1990 

Achrysophagus sp. Turkey Soylu & Urel 1977 

Anagyrus bohemani Spain 

India 

Carrero 1980a 

Varma 1977 

Anagyrus greeni Indonesia Kalshoven 1981 

Anagyrus pseudococci Italy 

Turkey 

USA 

USSR 

Viggiani 1975a 



Niyazov 1969 

Anagyrus sp. nr sawadai USA Meyerdirk et al. 1982 

Blepyrus insularis India Chacko et al. 1977 

Blepyrus saccharicola California Bennett & Hughes 1959 

Chrysoplatycerus splendens USA Bartlett & Lloyd 1958; 

Summy et al. 1986 

Clausenia josefi Mediterranean Niyazov 1969 

Coccidoxenoides (= Pauridia) peregrinus China 

India 

Italy 

USA 

Bartlett 1978 

Krishnamoorthy & Mani 1989a; Mani 1994 

Viggiani & Maresca 1973 

Meyerdirk et al. 1978, 1979, 1982 



Species 

HYMENOPTERA 



Leptomastidea abnormis Spain 

Turkey 

USA 

Leptomastix dactylopii Brazil 

India 

USA 

Carrero 1980a 



Mani 1995 

Mani 1995 


Leptomastix nigrocoxalis India Prakasan & Kumar 1985 

Leptomastix trilongifasciatus India Kalshoven 1981 

Ophelosia crawfordi Bartlett & Lloyd 1958 

Pseudaphycus angelicus California Bartlett & Lloyd 1958 

Pseudaphycus maculipennis USSR Sinadskii & Kozarzhevskaya 1980 

Pseudaphycus perdignus Bennett & Hughes 1959 

Sympherobius barberi USA Meyerdirk et al. 1982 

Timberlakia gilva South Africa Prinsloo 1982 


Allotropa citri China Bartlett & Lloyd 1958 

Allotropa kamburovi South Africa Annecke & Prinsloo 1977 

Allotropa mecrida USSR Niyazov 1969 

4.15 Planococcus citri 295


The citrus mealybug is also attacked in most regions by encyrtid and 

sometimes by platygasterid parasitoids (Table 4.15.2), several of which are 

capable of having a significant effect in warm climates. If the origin of 

P. citri is really China, it is surprising that there are not reports of a number 

of specific or near specific parasitoids from that region. Indeed the species 

most commonly employed for biological control is Leptomastix dactylopii 

which is believed to be native to Brazil. 

A few fungi attack P. citri under humid conditions (Table 4.15.3). 


Any account of the biological control of P. citri is complicated by the facts that 

(i) it has been confused with other species on a number of occasions, so that 

early records are often unreliable, (ii) documentation of some introductions is 

poor or lacking, and (iii) natural enemies (that also attack it) have often been 

introduced in programs aimed at other mealybugs. Table 4.15.4 summarises 

the main introductions for biological control of P. citri. 

CALIFORNIA 

It is convenient to outline, first, the prolonged attempts against P. citri in 

California, since programs elsewhere almost always draw extensively on 

experience there. Furthermore, since the first introduction of Cryptolaemus 

montrouzieri from Australia to California in 1891Ð92, there have been few 

parasites or predators used in the control of any economically important 

mealybug anywhere in the world that have not also been tested against 

P. citri in California, in the hope that they might attack it also (Bartlett 

1978). 

C. montrouzieri was mass reared and released in California against 

P. citri with some success, but repeated releases were required to achieve 

satisfactory control. Another coccinellid Nephus (= Scymnus) bipunctatus 

(under the name of Cryptogonus orbiculus) was introduced in 1910 from the 

Philippines, but did not become established (Bartlett 1978). However, 

establishment followed the introduction from Sicily in 1914 of the parasitoid 

Leptomastidea abnormis, although control was only partly successful 

(Viereck 1915; Smith 1917). The coccinellid Scymnus binaevatus from 

South Africa was established in 1921, but persists only in small numbers. 

Unsuccessful attempts were made to establish the encyrtid 

Coccidoxenoides peregrinus from Hawaii where it was having a major 

impact on P. citri, misidentified at the time as Planococcus kraunhiae. 

However, progeny of a single female from South China in 1950 allowed the 

species to become established, although at a low level (Flanders 1957).

Table 4.15.2 Parasitoids of Planococcus citri 

Species 

HYMENOPTERA 

ENCYRTIDAE 


Achrysophagus sp. Turkey Soylu & Urel 1977 

Anagyrus bohemani Spain 

India 

Carrero 1980a 

Varma 1977 

Anagyrus greeni Indonesia Kalshoven 1981 

Anagyrus pseudococci Italy 

Turkey 

USA 

USSR 




Niyazov 1969 

Anagyrus sp. nr sawadai USA Meyerdirk et al. 1982 

Blepyrus insularis India Chacko et al. 1977 

Blepyrus saccharicola California Bennett & Hughes 1959 

Chrysoplatycerus splendens USA Bartlett & Lloyd 1958; 

Summy et al. 1986 

Clausenia josefi Mediterranean Niyazov 1969 

Coccidoxenoides (= Pauridia) peregrinus China 

India 

Italy 

USA 

Leptomastidea abnormis Spain 

Turkey 

USA 


India 

USA 

Bartlett 1978 

Krishnamoorthy & Mani 1989a; Mani 1994 

Viggiani & Maresca 1973 


Carrero 1980a 



Mani 1995 

Mani 1995 


Leptomastix nigrocoxalis India Prakasan & Kumar 1985 


Table 4.15.2 (contÕd) Parasitoids of Planococcus citri 

Species 

HYMENOPTERA 



Leptomastix trilongifasciatus India Kalshoven 1981 

Ophelosia crawfordi Bartlett & Lloyd 1958 

Pseudaphycus angelicus California Bartlett & Lloyd 1958 

Pseudaphycus maculipennis USSR Sinadskii & Kozarzhevskaya 1980 

Pseudaphycus perdignus Bennett & Hughes 1959 

Sympherobius barberi USA Meyerdirk et al. 1982 

Timberlakia gilva South Africa Prinsloo 1982 


Allotropa citri China Bartlett & Lloyd 1958 

Allotropa kamburovi South Africa Annecke & Prinsloo 1977 

Allotropa mecrida USSR Niyazov 1969 

Table 4.15.3 Pathogens attacking Planococcus citri 

Species Country Reference 

Aspergillus flavus Cuba Martinez & Bravo 1989 

Cladosporium oxysporum South Africa Samways 1983; Samways & Grech 1986 

Entomophthora fresenii Indonesia Kalshoven 1981 

Entomophthora fumosa Australia Murray 1978; Samal et al. 1978 

Fusarium sp. Easter Is Ripa et al. 1995 


4.15 Planococcus citri 299 

The encyrtid Leptomastix dactylopii, introduced in 1934 from Brazil, 

was mass reared and released. Recoveries were made over a number of years 

following each spring or summer release, but not following the ensuing 

winter. Eventually, a few managed to overwinter, resulting in a low level 

population (Compere 1939). 

The South China platygasterid, Allotropa citri, introduced in 1950, 

attacked 1st and 2nd instar P. citri, but mass releases over some 6 years 

resulted in few field recoveries and it is not thought to be established 

(Bartlett 1978). 

A somewhat polyphagous encyrtid, Anagyrus pseudococci, was 

unsuccessfully introduced from Brazil in 1934 and 1953. The same or a 

similar species, Anagyrus sp. nr pseudococci, was brought in from Italy in 

1955, but was established only briefly (Bartlett 1978). 

The coccinellid Exochomus metallicus from Eritrea was established 

from introductions in 1954 against citricola scale (Coccus pseudomagnoliarum) 

and black scales and attacks P. citri on plants other than citrus 

(Bartlett 1978). 

In spite of this series of introductions, the natural enemies of P. citri in 

California do not, unaided, maintain populations at sub-economic levels, 

mainly because, it is claimed, climatic conditions permit the overwintering 

of, at best, inadequate populations. Cryptolaemus montrouzieri has often 

provided spectacular control of high populations, but is unable to maintain 

its numbers on low prey populations and disappears, requiring 

reintroduction. Methods are available for its low cost mass production 

(Fisher 1963). 

The encyrtid Leptomastidea abnormis is of considerable value in 

attacking young mealybugs sheltering under citrus fruit sepals in spring and 

autumn, but is adversely affected by high temperatures. Both Leptomastix 

dactylopii and Coccidoxenoides peregrinus build up high numbers 

following mass releases, but crash over winter and require repeated mass 

releases to maintain effectiveness. 

AUSTRALIA 

P. citri can be an important pest of citrus, but also attacks passionfruit and 

custard apple in warmer regions. The earliest attempts at biological control 

were in Western Australia, where Cryptolaemus montrouzieri was 

introduced from New South Wales in 1902 and an unidentified coccinellid 

from Spain in 1903. Only the former was established and it rapidly became 

an important factor in the successful control of mealybugs there (Wilson 

1960).


Before biological control was attempted in Queensland in 1980, six 

natural enemies were recorded, the coccinellids Cryptolaemus montrouzieri, 

and Harmonia octomaculata (= Coccinella arcuata) the chrysopids 

Chrysopa sp. and Oligochrysa lutea (all 4 native) and the exotic encyrtids 

Leptomastidea abnormis and Coccidoxenoides peregrinus. However, they 

were unable to maintain infestations consistently at acceptable commercial 

levels. Attack by a fungus similar to Entomophthora fumosa caused up to 

58.1% mortality of 3rd instar nymphs and adults on passionfruit during wet 

periods (Murray 1978, 1982; Smith et al. 1988). 

The Brazilian encyrtid Leptomastix dactylopii was imported from 

California and approximately 2.5 million adults released between 1980 and 

1987. It established readily and became the commonest natural enemy of 

P. citri throughout southeast Queensland, reducing mealybug infestations, 

averaging 38% of fruit in early December, to an acceptable level of 5% or 

less at harvest in April. Parasitoid numbers were lowest during winter and 

spring and augmentative releases of 5 to 10 thousand parasitoids per hectare 

in spring to early summer advanced parasitoid activity by 6 weeks. 

When no releases were made, the parasitoid was first recorded in early 

February and was present in an average of 55% of mealybug-infested fruit 

by mid-March. The mealybug infestation peaked at an average of 47% in 

mid-December but, by late April, only dropped to 10% and the presence of 

the mealybug on 25% or more fruit from December to March usually 

resulted in excessive amounts of sooty mould. C. montrouzieri was recorded 

on a maximum of 5% of mealybug-infested fruit, and augmentative release 

failed to increase this level. Augmentative release of L. dactylopii was found 

to be at least as cheap as pesticides and far more compatible with IPM of 

other citrus pests (Smith et al. 1988). 

Placing sticky bands around the trunks of custard apple trees reduced the 

numbers of the ant, Pheidole megacephala, and lowered, somewhat, the 

numbers of P. citri. Parasitisation of P. citri by Leptomastidea abnormis was 

low and unaffected by banding, but there were more predators (especially 

Oligochrysa lutea, Cryptolaemus montrouzieri and Syrphus sp.). 

Nevertheless, natural enemies were still unable to maintain P. citri at 

acceptable levels (Murray 1982). 

BERMUDA 

Seven species of parasitoid and four predators were introduced between 

1951 and 1955, mainly from California, but originating elsewhere: 

Coccidoxenoides peregrinus from Hawaii and south China, Leptomastix 

dactylopii (South American race) Leptomastidea abnormis, Pseudaphycus 

perdignus, Anagyrus pseudococci, Blephyrus saccharicola and Allotropa 

citri. Only C. peregrinus, L. dactylopii and L. abnormis were established

BRAZIL 

CHILE 

CHINA 

CUBA 

CYPRUS 

FRANCE 


and, of these, L. dactylopii may have been the local form. Of the predators, 

Cryptolaemus montrouzieri was established, but only briefly and two 

species of Hyperaspis and Scymnus sordidus failed to breed on P. citri 

(Simmonds 1957; Bennett and Hughes 1959). 

P. citri is common on a range of plants, but citrus is seldom infested. In 

1939 Leptomastidea abnormis and Leptomastix dactylopii were 

commonly reared from it, and it was attacked by numerous predators, 

including Hyperaspis c-nigrum, Nephus sp., Diomus sp., lacewings and 

cecidomyiids. Anagyrus pseudococci was not recorded, although it was 

present in Argentina (Compere 1939). 

Cryptolaemus montrouzieri was introduced in 1931, 1933 and 1939, but 

establishment is not recorded, although it is present on Easter Is (Ripa et al. 

1995). Extensive releases of two parasitoids from California resulted in 

establishment: Leptomastidea abnormis during 1931Ð36 and Leptomastix 

dactylopii in 1936 and 1938: both of these and Coccidoxenoides peregrinus 

were also established on Easter Is (Ripa et al. 1995). Attempts failed in 1954 

to establish from California: Allotropa citri, Anagyrus pseudococci, 

Coccidoxenoides peregrinus and Pseudaphycus perdignus (Duran 1944; 

Gonzalez and Rojas 1966). 

Four species of parasitic wasp were introduced from China to California in 

1950: a uniparental species of Coccidoxenoides and a biparental species of 

Allotropa from Guangzhou, a biparental species of Pseudaphycus from 

Taiwan and a biparental species of Coccophagus from Hong Kong (Flanders 

1951). 

Seven natural enemies attack P. citri on coffee including a Leptomastix sp.. 

A cecidomyiid predator was commonest (Martinez et al. 1992). A fungus 

Aspergillus flavus was also detected (Martinez and Bravo 1989). 

Leptomastix dactylopii was introduced from Italy in 1977 and became 

established, attaining a parasitisation rate of 15% in 1979. At harvest there 

were far fewer P. citrus on the fruit than in a plot that had received 3 

applications of insecticide (Krambias and Kontzonis 1980). 

Cryptolaemus montrouzieri was introduced from California in 1918 and 

became established, but overwinter survival was low (Marchal 1921, 1922; 

Poutiers 1922; Marchal and Pussard 1938). Of the 6 species of mealybug on 

citrus in the Mediterranean basin, P. citri is the most injurious. Damage by it 

in France is such that an economic injury level of 2% infested fruit has been


GREECE 

INDIA 

set. Mass rearing and release of Leptomastix dactylopii, which requires the 

use of fewer mealybugs for laboratory rearing than the coccinellid 

C. montruzieri and is cheaper, is preferred to that of the coccinellid if only 

one natural enemy is to be used. However, it is preferable to employ both, 

with C. moutrouzieri destroying high populations of females and eggs and 

L. dactylopii parasitising nymphs, even if populations are scattered (Panis 

1977, 1979). 

Cryptolaemus montrouzieri was liberated against P. citri on potted orange 

trees in a glasshouse at 25 to 30¡C and 55 to 70% RH and the results compared 

with the release of Nephus (= Scymnus) reunioni or the application of 

insecticide. C. montrouzieri significantly reduced mealybug populations 

and was as effective as treatment with methidathion (Hamid and Michelakis 

1994). 

Leptomastix dactylopii was introduced in 1983 to Bangalore and rapidly 

became established on P. citri on mandarins and coffee, causing up to 100% 

parasitisation (Nagarkatti et al. 1992). Seven years later in 1991, P. citri was 

being attacked on lemon and lime by L. dactylopii and the more abundant 

indigenous Coccidoxenoides peregrinus, which was causing 10 to 30% 

parasitisation. C. peregrinus attacks preferentially the early and Leptomastix 

dactylopii the later instars (Krishnamoorthy and Mani 1989a; Mani 1994). 

In 1984 L. dactylopii was released in a lime orchard in Karnataka. Prior to 

release, infestation by P. citri ranged from 38 to 65%, but establishment of 

the parasitoid led to excellent control within 4 months and no insecticides 

were required in following seasons. The parasitoid was shown to have 

migrated about 0.5 km in a 2-year period and a mean of 2.3 adult parasitoids 

were recovered from each infested fruit in an orange orchard 

(Krishnamoorthy 1990). Parasitisation of P. citri and other mealybugs on 

coffee ranged, in different years, between 19 and 45% (Reddy et al. 1988) 

and 0 and 85% (Reddy et al. 1992). Although L. dactylopii greatly reduced 

the population of P. citri, augmentative releases were required for 

continuing control in a coffee plantation (Reddy and Bhat 1993). 

Native predators of P. citri on coffee include the lycaenid, Spalgis epius, 

the coccinellid, Pullus pallidicollis, and the cecidomyiid Triommata 

coccidivora (Prakasan 1987). On citrus the chrysopids Mallada boninensis, 

Odontochrysa (= Chrysopa) lacciperda, Mallada basalis and Chrysoperla 

carnea were found (Krishnamoorthy and Mani 1989b); and on guava 

Odontochrysa lacciperda, Spalgis epius and Cryptolaemus montrouzieri 

(Mani and Krishnamoorthy 1990).

ISRAEL 


Releases of Cryptolaemus montrouzieri on a coffee estate in Kerala 

virtually eliminated P. citri, but the coccinellid could not be found thereafter 

for some 6 months, when it reappeared about 10 km distant and virtually 

eliminated mealybugs from an infestation (Chacko et al. 1978). The 

requirement that each female consume at least 8 P. citri for normal egg 

production (192 eggs/female) was suggested as the reason for the poor 

establishment of C. montrouzieri when only low populations of P. citri are 

available (Reddy et al. 1991). 

The citrus mealybug is a serious pest of citrus and ornamentals of tropical 

origin. It develops on the fruit and roots of young trees, the main damage 

being caused by individuals settling beneath the sepals of citrus fruit and 

injuring the fruit: honeydew produced also attracts fruit-piercing moths 

(Mendel et al. 1992). 

Two native encyrtids, Anagyrus pseudococci and Leptomastidea 

abnormis, attack P. citri which is, nevertheless, an important pest of citrus 

and some other crops (Rosen and Ršssler 1966). 

Unsuccessful attempts were made to establish Cryptolaemus 

montrouzieri in 1924 from Egypt (Bodenheimer and Guttfeld 1929; Mason 

1941) and 1958 (Rosen 1967a), but a Spanish strain introduced in the 1980s 

finally established (Mendel et al. 1992). Leptomastix dactylopii imported 

from Canada in 1941 was only briefly established (Rivnay 1960). Clausenia 

purpurea, a parasitoid of Pseudococcus citriculus introduced from Japan in 

1940, attacks P. citri, but has little effect on its density (Rosen 1964). 

P. citrus became a serious pest on recently established grapefruit in 

southern Negev. The encyrtid Anagyrus pseudococci was present in very 

low numbers and Chrysoperla carnea was active, but only in spring. The 

coccinellid Chilocorus bipustulatus was more abundant and was considered 

to have potential as a biological control agent (Berlinger et al. 1979). Seven 

parasitoids and 10 predators have been recorded, of which Anagyrus 

pseudococci (Encyrtidae) and Sympherobius sanctus (Chrysopidae) were 

found in significant numbers by Mendel et al. (1992). However, 

development of A. pseudococci is restricted by winter temperatures. Its 

development threshold is 13¡C and no eggs are laid below 15¡C, whereas 

P. citri still lays eggs at 13¡C and its development threshold is 8.4¡C. By the 

time in late spring when the population of A. pseudococci starts to built up, 

P. citri has already settled under the sepals, where it is well protected from 

the parasitoid. Climatic conditions were thus regarded as unfavourable for 

existing natural enemies (Mendel et al. 1992).


ITALY 

Infestations of P. citri develop on citrus in Sicily, Procida, Sardinia and parts 

of mainland Italy, especially where it is protected by ants from its native 

parasitoids, Leptomastidea abnormis and Anagyrus pseudococci (Zinna 

1960). Cryptolaemus montrouzieri has been imported a number of times 

since 1908 and has become established in some warmer areas, but is so 

reduced in numbers during winter that satisfactory control is not obtained 

without supplementations (Constantino 1935; Liotta 1965; Liotta and Mineo 

1965). Coccidoxenoides peregrinus and Leptomastix dactylopii were 

introduced to Procida in 1956Ð57, but did not survive the winter (Bartlett 

1978). L. dactylopii was reintroduced in 1974 to this island, to the mainland 

(Campania, Calabria), and to Sardinia and Sicily. At almost all 15 release 

sites it afforded, initially, a high level of parasitisation (Viggiani 1975a,b). 

Small overwintering populations, reduced further by hyperparasitoids and 

fungus diseases, persisted in some areas, but required supplementation for 

control (Mineo and Viggiani 1975a,b). L. dactylopii was again released in 

1979 on Sicily and gave good results (5% of fruit infested), but it is unclear 

whether or not it is able to overwinter (Barbagallo et al. 1982; Longo and 

Benfatto 1982). Release of L. dactylopii in a mainland orange orchard in 

Calabria led to a reduction in infested fruit from 80.9% to 12% and only 

7.3% of the fruit was unmarketable (Luppino 1979). It now seems that 

L. dactylopii and Cryptolaemus montrouzieri are mass reared and released 

each year against P. citri (Raciti et al. 1995). 

Insecticides are seldom required against P. citri in Sardinia, where 

L. dactylopii may cause 96% cumulative parasitisation and where 

Cryptolaemus montrouzieri and Nephus (= Scymnus) reunioni may increase 

to more than 100 individuals per tree (Ortu 1982; Ortu and Prota 1983). 


Mealybugs, including P. citri, were causing up to 75% reduction of coffee 

yield in the highlands near Wau in the mid fifties. Introduction of 

Cryptolaemus affinis in 1957 from the lower Markham Valley resulted in its 

rapid spread and a substantial reduction of the infestations within one season 

(Szent-Ivany 1963). 

PERU 

Very good control of P. citri is given in some areas by the encyrtid 

Coccidoxenoides peregrinus, which is restricted to the citrus mealybug and 

appears to have arrived accidentally in 1963 (Salazar 1972). 

SOUTH AFRICA 

The history of introductions to South Africa against P. citri is confused 

because of misidentifications of other mealybugs for this species. However, 

Bedford (1976) reports that P. citri is under biological control.


When the ant Anoplolepis custodiens was excluded from guava trees 

bearing P. citri at Nelspruit, the population of both ants and mealybugs 

dropped to half. Without ants, the mealybugs were heavily preyed upon by 

the ant-intolerant Exochomus flavipes and the ant-tolerant Scymnus spp. 

Later, the mealybugs were almost eliminated by the fungus Cladosporium 

sp. nr oxysporum (Samways 1983). 

SPAIN 

P. citri is parasitised by two indigenous encyrtids, Leptomastidea abnormis 

and Anagyrus bohemani, but at a low level (Carrero 1980a). Cryptolaemus 

montrouzieri was introduced and established before 1928 and produces 

control in the warmer months (Gomez 1951; Carrero 1980b). Leptomastix 

dactylopii was introduced in 1948 from California (Gomez 1951) and in 

1977 from Italy, but did not become established (Carrero 1980b). 

USA (OTHER THAN CALIFORNIA) 

Florida 

Cryptolaemus montrouzieri was introduced to Florida in 1930 for the control 

of P. citri on citrus and bulbs. It became established but failed to overwinter 

in sufficient numbers to achieve adequate control (Bartlett 1978; Muma 

1954, Watson 1932). 

Hawaii 

P. citri (originally misidentified as P. kraunhiae) was the target of many 

introductions (Swezey 1931), although not as severe a pest as on the 

mainland. Cryptolaemus montrouzieri was introduced from Australia in 

1894, Leptomastidea abnormis from California in 1915 and Leptomastix 

dactylopii also from California in 1946. All became established. 

Texas 

In 1970, Coccidoxenoides peregrinus was the dominant parasitoid of 3 

present on P. citri attacking grapefruit and Sympherobius barberi the 

commonest predator (Dean et al. 1971). In 1977 Leptomastix dactylopii, 

which had been introduced from California in 1970, and Anagyrus sp. nr 

sawadai were found for the first time on P. citri in Texas. L. dactylopii was 

the most abundant parasite in mid-August (parasitising 20.7% of P. citri), 

the reputedly indigenous Coccidoxenoides peregrinus (with 48.5%) the 

most abundant in late August, whereas Anagyrus sp. (with 4.3%) was the 

only parasitoid recovered in mid-September. A hyperparasitoid 

Prochiloneurus dactylopii attacked 1% of the primary parasitoids in mid- 

August (Meyerdirk et al. 1978). Release of 5 encyrtid parasitoids on 

glasshouse citrus resulted in the rapid suppression of P. citri. Leptomastidea 

abnormis, Anagyrus pseudococci and Leptomastix dactylopii persisted for 

periods between 24 and 32 weeks and maintained the host at low densities, 

whereas Chrysoplatycerus splendens and Coccidoxenoides peregrinus


USSR 

persisted for only 20 weeks (Summy et al. 1986). The coccinellid Diomus 

pumilio, whose biology is described, was introduced from South Australia 

and is a potentially valuable predator (Meyerdirk 1983). 

P. citri can be a serious pest of grape vines, citrus, fig and pomegranate. 

Anagyrus pseudococci from Surkham Dalya and Leptomastix dactylopii and 

Leptomastidea abnormis from California were introduced to Uzbekistan 

commencing in 1959 and resulted in establishment (Roxanova and Loseva 

1963). Anagyrus pseudococci destroys up to 75% of P. citri in areas not 

treated with insecticides in the south of European Russia and in Soviet 

Central Asia. The next most important parasitoid, Allotropa mecrida 

attacked up to 20% in Turkmenia in 1967 and in Georgia. In 1960, 

Leptomastidea abnormis and Leptomastix dactylopii were introduced from 

USA into Georgia and Turkmenia. In Transcaucasia and Soviet Central Asia 

the hyperparasitoid Thysanus subaeneus attacks 18 to 20% of Allotropa 

mecrida. Other hyperparasitoids are Pachyneuron solitarius and 

Neoprochiloneurus bolivari. 

One of the most effective predators of P. citri is Cryptolaemus 

montrouzieri, introduced from Egypt in 1932 to the Black Sea area. Others 

are Coccinella septempunctata, Hyperaspis polita, Nephus bipunctatus, 

Scymnus apetzi, S. subvillosus, and S. biguttatus which were recorded in 

Turkmenia. The larvae of the fly Leucopis alticeps and of the lacewing 

Chrysoperla carnea are able to devastate all stages of P. citri. The 

coccinellids were parasitised by Homalotylus sp. and the lacewing by 

Telenomus acrobates (Niyazov 1969). 

The coccinellid Nephus reunioni was introduced into southern areas in 

1978 and has reduced P. citri on grape vines. It is capable of overwintering, 

but with high mortality, and is more tolerant of moisture conditions than 

Cryptolaemus montrouzieri (Orlinskii et al. 1989).

Table 4.15.4 Introductions for the biological control of Planococcus citri 

Species 

HYMENOPTERA 

ENCYRTIDAE 


Anagyrus kivuensis Kenya California 1948 Ð Bartlett & Lloyd 1958 

Anagyrus pseudococci Brazil 

California 

California 

California 

Bermuda 

Chile 

1934 

1953 

1951Ð54 

1954 

Ð 

+ 

Ð 

Ð 

Bartlett & Lloyd 1958 



Gonzalez & Rojas 1966 

Anagyrus sp. nr pseudococci Italy California 1955, 1965 Ð Bartlett 1978, 


Anagyrus sp. nr sawadai Texas + Meyerdirk et al. 1978 

Blepyrus saccharicola California Bermuda 1951Ð54 Ð Bennett & Hughes 1959 

Coccidoxenoides peregrinus Hawaii California Ð Armitage 1920 

China California + Flanders 1951 

China Bermuda 1951Ð54 + Bennett & Hughes 1959 

California Chile 1954 Ð Gonzalez & Rojas 1966 

Peru 1963 + Salazar 1972 

Leptomastidea abnormis Sicily 

California 

California 

USA 

California 

Chile 

Bermuda 

USSR 

1914 

1931 

1951Ð54 

1960 

+ 

+ 

+ 

Viereck 1915; Smith 1917 



Niyazov 1969 

Leptomastidea sp. nr abnormis Mexico California 1956 Ð Bartlett & Lloyd 1958 


Table 4.15.4 (contÕd) Introductions for the biological control of Planococcus citri 

Species 

HYMENOPTERA 




California 

California 

California 

California 

Italy 

California 

Chile 

Chile 

Bermuda 

Turkey 

Texas 

Australia 

India 

India 

Israel 

Italy 

Cyprus 

Sardinia 

1934 

1936 

1958 

1951Ð54 

1970 

1980 

1983Ð85 

1984 

1956Ð7 

1977 

1974 

+ 

Ð 

+ 

? 

+ 

+ 

+ 

+ 

+ 

Ð 

Ð (early) 

+ (later) 

+ 

Ð 

Compere 1939 




Tuncyurek 1970 


Smith et al. 1988 

Nagarkatti et al. 1992 

Krishnamoorthy 1990, 

Prakasan & Bhat 1985; 

Krishnamoorthy & Singh 1987; 

Prakasan 1987; Reddy et al. 1988; 

Mani 1994 

Rivnay 1960 

Luppino 1979; Longo & Benfatto 

1982 

Viggiani 1975a,b; Mineo & 


Krambias & Kontzonis 1980 

Viggiani 1975a,b; Delrio et al. 

1981; Ortu & Prota 1981; 

Ortu 1982 

Sicily 1979 Ð Barbagallo et al. 1982 

Longo & Benfatto 1982 

Viggiani 1975a,b 

Italy Spain 1977 Ð Carrero 1980b 

USA USSR 1960 Niyazov 1969 



Species 

HYMENOPTERA 



Pseudaphycus perdignus Eritrea California 1953 Ð Bartlett & Lloyd 1958 

California Bermuda 1951Ð54 Ð Bennett & Hughes 1959 

Tropidophryne melvillei Kenya California 1948 Ð Bartlett & Lloyd 1958 

HYMENOPTERA 


Allotropa citri China California ? Flanders 1951; Bartlett & Lloyd 

1958 

California Bermuda Ð Bennett & Hughes 1959 

California Chile 1954 Ð Gonzalez & Rojas 1966 

COLEOPTERA 

COCCINELLIDAE 

Brumus suturalis India California 1955 Ð Bartlett & Lloyd 1958 

Chilocorus angolensis Kenya California 1948 Ð Bartlett & Lloyd 1958 

Cryptolaemus montrouzieri Greece Ð Argyriou 1970 

Sardinia


Species 

COLEOPTERA 



California Bermuda 1955 + Bennett & Hughes 1959 

California Chile 

India 

Indonesia 

1931 

1933 

1934 

Ð 

Ð 

? 

+ 

Ð 




Prakasan 1987 


Spain Israel 1980s + Mendel et al. 1992 

Diomus pumilio (= D. flavifrons) Australia Texas + Meyerdirk 1983 

Exochomus flavipes Kenya California 1948 Ð Bartlett & Lloyd 1958 

Exochomus metallicus Eritrea California 1954 + Bartlett 1978 

Hyperaspis jucunda Trinidad California 1955 Ð Bartlett & Lloyd 1958 

Hyperaspis sp. nr globula Mexico California 1954 Ð Bartlett & Lloyd 1958 

Hyperaspis sp. Eritrea California 1953 Ð Bartlett & Lloyd 1958 

Hyperaspis 2 ´ spp. California Bermuda 1958 Ð Bennett & Hughes 1959 

Nephus (=Scymnus) bipunctatus Philippines California 1910 Ð Bartlett 1978 

Nephus (=Scymnus) reunioni East Africa USSR 1978 + Ershova & Orlinskii 1982; Orlinskii 

et al. 1989 

Nephus sp. Trinidad California 1958 Ð Bartlett & Lloyd 1958 

Platynaspis (?) sp. Eritrea California 1953 Ð Bartlett & Lloyd 1958 

Scymnus binaevatus South Africa California + Smith 1923 

Scymnus quadrivittatus Kenya California 1948 Ð Bartlett & Lloyd 1958 

Scymnus sordidus California Bermuda 1955 Ð Bennett & Hughes 1959 


Biology of important natural enemies 


Anagyrus pseudococci Hym.: Encyrtidae 

This is a solitary endoparasite of 2nd, 3rd and 4th instar mealybugs, but 

prefers 3rd instar and egg-laying females. It is believed to be native to the 

Mediterranean. A. pseudococci has been known since 1913 in Italy as a 

widespread parasitoid of P. citri and, in Israel, attacking both P. citri and 

Pseudococcus citriculus (Rivnay 1960; Rosen 1964). It is common on 

P. citri in Argentina (Compere 1939). In the laboratory it develops 

successfully on Pseudococcus fragilis, P. longispinus and P. obscurus. 

Females lay about 45 eggs at the rate of 3 to 4 per day. In the laboratory 

these hatch in 44 hours at 27¡C and the life cycle takes 17 to 18 days (Bartlett 

1978) or 15.5 days at 25.6¡C and 60% RH for a Californian strain, which 

also had a life span of 8.2 days for virgin and 6.9 days for mated females. 

Virgin females produce males. A. pseudococci is most active in the spring 

and autumn (Domenichini 1952; Avidov et al. 1967; Rivnay 1968; Chandler 

et al. 1980). 

Progeny production increased and longevity decreased with increase in 

temperature between 18¡ and 30¡C (Tingle and Copland 1989). Most 

progeny are produced between 27¡ and 30¡C and the threshold for 

development is 13.06¡C for males and 12.57¡C for females (Islam and Jahan 

1993). In the laboratory maximum egg production was achieved when 50% 

honey solution was provided (Islam and Jahan 1992, 1993). Oviposition 

behaviour is described by Islam (1992). About 40% of parasitoid eggs laid in 

P. citri may be lost due to encapsulation (Blumberg et al. 1995). In 

Argentina, larvae of A. pseudococci are attacked by the hyperparasitoid 

Coccophagus heteropneusticus (Compere 1939). 

Coccidoxenoides peregrinus Hym.: Encyrtidae 

This parasitoid, earlier widely known as Pauridia peregrina, is probably 

native to southern China (Bartlett 1978), although Meyerdirk et al. (1978) 

suggest that it is indigenous to Texas. It has been reported, inter alia, from 

India, Japan, Philippines, Fiji, Hawaii and Uganda. It is a solitary 

endoparasitoid of 1st, 2nd and 3rd instar female Planococcus citri and 

P. kenyae and 1st and 2nd instar males. It is normally parthenogenetic, but 

there are rare males. In Uganda it parasitises P. kenyae (Armitage 1920; 

Essig 1931) and, in Peru P. citri (Beingolea 1969). In India it completed its 

development only in P. citri, although it attacked other mealybugs 

(Krishnamoorthy and Mani 1989a). 

Females commence oviposition shortly after emergence, and continue 

for about 2 days. At 27¡C larval development takes 11 to 12 days and the 

pupal stage 16 to 18 days (Zinna 1960; Fisher 1963). However, in India,


development took 23 to 27 days and adults survived 4 to 9 days at 28 ± 2¡C 

(Krishnamoorthy and Mani 1989a). In 1991 in Karnataka, C. peregrinus was 

more abundant than Leptomastix dactylopii on P. citri on lemon and acid 

lime and was responsible for the decline of mealybug populations (Mani 

1994). 

C. peregrinus has been introduced into California (Flanders 1951), Italy 

(Bartlett 1978) and Bermuda (Bennett and Hughes 1959). 

Cryptolaemus montrouzieri Col.: Coccinellidae 

This general predator of mealybugs, which also feeds on some other scales 

(Eriococcus sp., Pulvinaria spp.) and aphids, is native to eastern Australia. It 

is the most widely distributed of all natural enemies of mealybugs, a count in 

1978, covering the past 80 years, listing more than 40 countries, geographic 

areas or islands into which it has been imported. In many instances it was 

introduced against mealybugs other than P. citri and sometimes against 

coccids such as Pulvinaria spp., which produce egg masses similar to those 

of mealybugs (Bartlett 1978). 

Both larvae and adults feed voraciously on all mealybug stages, for 

example a larva is recorded as consuming an average of 3331 host eggs 

(Oncuer & Bayhan 1982) and females need to consume at least 8 P. citri for 

normal egg production (Reddy et al. 1991). C. montrouzieri does not 

distinguish between unparasitised P. citri and mealybugs parasitised by 

Leptomastix dactylopii (Prakasan and Bhat 1985). Adults mate 1 or 2 days 

after emergence and, 5 to 6 days later, females begin ovipositing in or near 

host egg masses. About 100 eggs are deposited in 1 month. These hatch in 4 

to 8 days, and wax-covered larvae develop in 12 to 20 days, so that the life 

cycle can be completed in slightly less than a month (27.7 days at 

25.5¡ ± 1¡C: Oncuer and Bayhan 1982), although there are usually only 4 

generations a year. Development stops below 10¡C and freezing 

temperatures are lethal. Pupae, and occasionally adults, are capable of 

hibernating. Hot dry climates are tolerated, but high humidities are said to be 

detrimental. C. montrouzieri thrives when host density is high and, under 

these conditions, is capable of providing spectacular control. However its 

searching ability and natural spread is poor, so it often dies out locally when 

hosts become scarce (Bodenheimer 1928; Cole 1933; Mineo 1967). 

Methods have been developed for the production of mealybugs and 

C. montrouzieri that permit the production and release of large numbers of 

the predators at low cost (Branigan 1916, Smith and Armitage 1920, 1931; 

Fisher 1963; Chacko et al. 1978; Oncuer and Koldas 1981).


Diomus pumilio Col.: Coccinellidae 

Details of the biology and voraciousness of this predator, which was 

introduced from South Australia to Texas for biological control of P. citri, 

are given by Meyerdirk (1983). 

Leptomastidea abnormis Hym.: Encyrtidae 

This solitary endoparasite is possibly native to the Mediterranean where it 

was first recognised attacking P. citri (Viereck 1915), although it is now 

widespread, occurring in eastern USA, Canada, Brazil (Compere 1939) and 

many other countries. 

L. abnormis strongly prefers small 2nd instar mealybugs for oviposition, 

but also attacks first and third instars. Females begin to search for hosts soon 

after emergence. The number of eggs laid varies from 57 to over 300, 

although it is reported that only about 33 survive to the adult stage. Fertilised 

eggs give rise to females and unfertilised eggs to males. 

The inconspicuously stalked eggs are laid free in the haemolymph and 

hatch in 36 to 72 hours. The larvae consume haemolymph at first but, in the 

last instar, consume the entire body contents. The tailed larvae complete 

development in about 8 days and the life cycle may be as short as 17 days in 

the laboratory at 26¡C (or 25 days at 25¡ to 27¡C and 50Ð70% RH). In the 

laboratory females attained their maximum progeny production at 24¡C and 

this remained constant up to 34¡C (Tingle and Copland 1989). In the field a 

generation in summer takes about 1 month. There may be 5 or 6 generations 

a year, adults living 11 days if provided with honey and water (Viereck 

1915; Smith 1916, 1917; Perez 1929; Rivnay and Perzelan 1943; Clancy 

1944; Viggiani and Maresca 1973). 

Leptomastix dactylopii Hym.: Encyrtidae 

This solitary endoparasitoid prefers 3rd instar and young (but not egglaying) 

females and occasionally attacks 1st and 2nd instars (Bartlett 1978; 

Mani 1995). It is presumed to be native to Brazil, although found also in the 

West Indies and parts of southern USA (Compere 1939). In the field it 

appears to be specific to P. citri (Bartlett 1978; Sinadskii and 

Kozarzhevskaya 1980; Nagarkatti et al. 1992), but it can be reared on 

Planococcus lilacinus (Mani 1995), P. pacificus (Nagarkatti et al. 1992), 

Phenacoccus solani (Lloyd 1964) and Pseudococcus comstocki (Clancy 

1944). Its reported attack on Dysmicoccus brevipes in Hawaii and on 

Pseudococcus vitis in USSR was probably on P. citri (Kobakhidze 1965; 

Bartlett 1978). 

It has been used in suppression of P. citri in USA (Fisher 1963), Procida 

island and mainland Italy (Luppino 1979), Cyprus (Krambias and Kontzonis 

1980) and India (Krishnamoorthy 1990).


Adults live up to 35 days and longer at 15¡ than at 7¡ or 25¡C (Yigit et al. 

1994) although maximum progeny are produced at 30¡ (Tingle and Copland 

1989). Parasitised hosts are generally rejected after simple antennal contact 

but, if not then, also following defensive behaviour of the host or possibly 

after detection of the egg stalk emerging from the surface of the host. If not 

rejected earlier, they may be rejected after insertion of the ovipositor 

(Baaren and Nenon 1994). About 18 eggs are laid each day, up to a total of 

300 per female. These hatch in 1.5 to 2 days at 28¡C and there are four larval 

instars, each of about 2 days. The pupal stage lasts 7 to 8 days. In Italy there 

are 6 (and a partial 7th) generations per year (Zinna 1959, 1960) and in 

Tashkent 5 generations (Roxanova and Loseva 1963). 

More males than females are produced from young than from old adult 

P. citri (Su and Li 1993; Mani 1995), more females from larger hosts and 

more males from smaller larval instars (Jong and Alphen 1988, 1989). 

Additional information on the biology of L. dactylopii is given by Lloyd 

(1958, 1964, 1966) and Tingle and Copland (1989). 

The original introduction of L. dactylopii from Brazil to California in 

1934 was based on a single pair (Compere 1939). The extent to which the 

progeny of this pair may have had genes from later introductions added to 

the gene pool is quite unclear. There may thus be good justification for 

obtaining fresh stock from matching climatic zones in Brazil if new 

introductions are to be made. 

Odontochrysa (= Chrysopa) lacciperda Neu.: Chrysopidae 

Details of the biology and voracity of this lacewing predator of P. citri are 

provided by Krishnamoorthy (1988). 

Pseudaphycus maculipennis Hym.: Encyrtidae 

This species, studied in Ukraine, for the biological control of P. citri, is said 

to be specific (Sinadskii and Kozarzkevskaya 1980). 

Scymnus includens Col.: Coccinellidae 

The life cycle and rearing details of this important predator of P. citri in Italy 

are described by Tranfaglia and Viggiani (1973). 

Spalgis epius Lep.: Lycaenidae 

The predatory larvae of this lycaenid butterfly are often the commonest 

natural enemies of P. citri in India. They also attack Planococcus lilacinus, 

Chloropulvinaria psidii and Ferrisia virgata (Chacko et al. 1977).

Comments 


P. citri is frequently only one of several pests of importance on the economic 

crops that it infests and, if biological control is contemplated, it is advisable 

to have its identity confirmed by a competent taxonomist. Its abundance is 

increased when it is tended by ants for its honeydew and often, by the 

injudicious use of insecticides (against it or accompanying pests). This is 

mainly because of the adverse effects on natural enemies and sometimes 

because low levels of insecticide may stimulate egg-laying. There has thus 

been considerable effort, within an Integrated Pest Management framework 

and with varying degrees of success, to develop biological control of each of 

the important pests in a complex, for example on citrus, grape vines and in 

glasshouses. This has also involved the careful selection of pesticides (if 

these are still required) that have the least possible adverse effect on the 

major natural enemies. 

Where natural enemies already present are not adequate, the almost 

universal response has been to introduce the encyrtid Leptomastix dactylopii 

and the coccinellid Cryptolaemus montrouzieri (if the latter is not already 

present as a result of introductions for other pests). 

Both species are affected by the winter in temperate zones and survive 

less well than P. citri in the Mediterranean region. Under these 

circumstances, classical biological control seldom provides economic 

control alone and requires augmentation of the natural enemies from time to 

time. 

Where P. citri is still a problem in the field in warm regions and 

Cryptolaemus montrouzieri is not present, serious thought should be given 

to introducing the latter. More importantly, however, if not already present 

Leptomastix dactylopii (of Brazilian origin) should be of highest priority, 

followed by Leptomastidea abnormis and Anagyrus pseudococci (both of 

Mediterranean origin and capable of maintaining populations of low levels 

under slightly cooler conditions). Under Southeast Asian and Pacific 

conditions, Coccidoxenoides peregrinus (of south China or Indian origin) 

deserves special attention. 

If Planococcus citri proves to be of south China origin (and this 

hypothesis requires confirmation) it is surprising that only two parasitoids 

have been reportedÑthe encyrtid Coccidoxenoides peregrinus and the 

platygasterid Allotropa citri. A thorough survey in this region might well 

reveal additional valuable specific or near specific parasitoids. There are 

good grounds for optimism that biological control of P. citri can be 

improved in warmer regions by establishing, if missing, any one of the 

foregoing natural enemies.


P. citri is one of a group of pests that commonly cause problems in 

glasshouses in Europe and North America. The mass production and release 

from time to time, almost always of a predator (especially Cryptolaemus 

montrouzieri, but sometimes also Nephus reunioni) and one or more 

encyrtid parasitoids (especially Leptomastix dactylopii and Leptomastidea 

abnormis but, on occasion also, Coccidoxenoides peregrinus, Anagyrus 

pseudococci and Chrysoplatycerus splendens) has generally removed the 

need to use insecticides. Examples of control in glasshouses include those 

from Belgium (Ronse 1990), Netherlands (Heanekam et al. 1987), France 

(Panis and Brun 1971), U.K. (Copland 1983, Hussey and Scopes 1985; 

Tingle and Copland 1988), Israel (Rubin 1985) and USA (Summy et al. 

1986).

4.16 Trichoplusia ni 

India 

20° 

Myanmar 

++ Laos 

0° 

20° 

China 

+ 

Thailand 

++ 

Cambodia 

++ 

Vietnam 

+ 

Malaysia 

Singapore 

Brunei 

P 

Indonesia 

Taiwan 

+ 

Philippines 

Australia 

Papua 

New Guinea 

317 

The cabbage looper, Trichoplusia ni, 

of North American origin, attacks cabbage (and other 

Brassicaceae), cotton, lettuce, tomatoes and a very wide range of other cultivated crops and 

wild hosts. In North America it is maintained for much of the time at sub-economic levels by a 

wide range of natural enemies, but damaging outbreaks do occur, particularly when its natural 

enemies are killed by insecticides applied against other pests in the same crop. 

The major predators, which together cause considerable mortality, are widely polyphagous, 

and hence are unlikely to be considered seriously as classical biological control agents. Several, 

among its 120 or so parasitoids are somewhat more host specific and are worth serious 

consideration. They include species of Trichogramma egg parasitoid; Copidosoma 

truncatellum (an egg-larval parasitoid); and the larval parasitoids Hyposoter exiguae, 

Microgaster brassicae and Voria ruralis. 

High larval mortality is frequently produced by a valuable, naturally occurring, nuclear 

polyhedral virus, particularly late in the season when T. ni populations are high and rainfall is 

adequate. 

There appear to be good reasons for optimism that the establishment of suitable missing 

natural enemies in regions into which T. ni has spread would assist in maintaining its populations 

at sub-economic levels. 

20° 

0° 

20°


Trichoplusia ni (HŸbner) 

Rating 

Origin 

Distribution 

Biology 

Lepidoptera: Noctuidae 

cabbage looper 

Synonym: In North America T. ni has sometimes been referred to 

as Autographa brassicae (Riley). 

Southeast Asia China 

7 ++ Myan, Thai, Camb + (all 14 southern Provinces) 

+ Viet 

P Indo 

T. ni is native to the southern half of North America. 

Widespread in southern Europe, North, East and South Africa, extending 

eastwards through Pakistan, India and Bangladesh to much of Southeast 

Asia, to China, Taiwan, Korea and Japan; not yet present in Papua New 

Guinea, Australia, New Zealand or the oceanic Pacific; present in South 

America in Argentina, Bolivia, Brazil, Chile, Colombia and Uruguay 

(Apablaza and Norero 1993; CIE 1974b). In North America, T. ni 

overwinters in the south, re-invading northern States each spring. 

Adult T. ni are mottled brownish in colour. The forewings, producing a span 

of about 3.8 cm, each bear an 8-shaped silvery mark near the middle. Adults 

are mostly active at night, but also on dull days. By day, they rest on the 

underside of host plants, in the debris at their base, or in vegetation bordering 

a cultivated crop. Adults are capable of flying long distancesÑ700 km 

northwards from southern Texas (Lingren et al. 1993) 161 km from land into 

the Gulf of Mexico, and up to 1500m in California (Kreasky et al. 1972). 

They move readily between cultivated and wild hosts. A female may lay 

her own bodyweight in eggs, but requires access to nectar and moisture to do 

so. After emerging from the pupa, there is a pre-ovipositional period of about 

4 days, after which mating begins and can occur up to 16 days. A female may 

produce well over 1000 viable eggs. Peak egg deposition is often correlated 

with the lunar cycle, a rapid rise in egg density on cotton occurring shortly

Host plants 

4.16 


319 

after full moon. There are 3 generations a year in southern California, but 

breeding is continuous in the Caribbean (McKinney 1944; Kishaba et al. 

1967; Ehler and van den Bosch 1974; Ehler 1977a; Debolt et al. 1984; 

Mitchell and Chalfant 1984). 

In cotton, a single egg is laid on the underside of a mature leaf in the 

upper half of the plant, but seldom in a terminal. On hatching, the larva 

generally feeds on the underside of the leaf near the egg, later moving from 

leaf to leaf as it passes through 5 instars during 2 to 4 weeks (Ehler 1977a). 

Total development time ranges from 19.9 days at 30¡C to 40.4 days at 20¡C 

(Jackson et al. 1969). In India at 25¡C the egg stage lasted 2.06 days, the 

larval stage of 5 instars 12.38 days, the prepupal 1 day, the pupal 7.27 days 

and the adult 7.32 days (Gaikwad et al. 1983). Additional data are provided 

by Chi and Tang (1993) and Yadav et al. (1983). Flight and mating activity 

are diminished at temperatures less than 16¡C and the threshold for larval 

development lies between 10¡C and 13¡C. The larva has 3 pairs of true legs 

on the thorax and 3 pairs of fleshy abdominal prolegs near the posterior end. 

It crawls by doubling up to form a loop, thus projecting the body forward. 

Larvae are green with a white lateral line and 2 whitish lines along the 

middle of the dorsal surface. After a brief prepupal period, pupation occurs 

in a loosely spun cocoon either on the underside of a leaf or in plant debris at 

the soil surface (Ehler 1977a). There is no diapause (Fye 1979). 

Eggs and larvae, but also pupae, of T. ni are believed to be readily 

transhipped in vegetables and cut flowers (Poe and Workman 1984). 

Male T. ni are powerfully attracted to the sex pheromone emitted by 

virgin females and will fly long distances upwind under its influence. The 

pheromone is produced in a gland situated dorsally between the 8th and 9th 

abdominal segments. Six components have been identified and are required 

to ensure specificity to T. ni. 

The major component is (Z)-7-dodecenyl 

acetate, known as looplure, which also attracts males of other looper species. 

Looplure, with or without other components, has been used for trapping 

males and also in mating disruption experiments. Male T. ni also produce a 

pheromone (with at least 3 components) which attracts both females 

(especially when starved) and males. ( Bjšstad et al. 1984; McLaughlin 

1984; Heath et al. 1992; Dunkelblum and Mazor 1993; Landolt 1995; 

Landolt et al. 1996). 

In 1966, larvae of T. ni were recorded causing damage to at least 119 species, 

varieties and cultivars in 29 families of plants (Sutherland 1966) and that 

number has increased steadily over the years to over 160 species in 36


Damage 

families, although cultivated brassicas are those most favoured when 

available (Martin et al. 1976a; Sutherland and Greene 1984). Brassicas and 

cotton are most frequently cited as being damaged, although the list of 

economic crops affected also includes asparagus, beans, sugarbeet, 

cantaloupes, capsicum, carrot, celery, maize (silks), cucumber, lettuce, 

parsley, pea, potato, soybean, spinach, squash, tobacco, tomato and 

watermelon. 

At times, serious infestations occur, but T. ni is generally regarded as a 

secondary pest whose numbers increase late in the season (Ehler 1977a,b). 

Differences in susceptibility to T. ni have been found in cabbage and 

related brassicas, in cotton and in lettuce (Cuthbert and Kishaba 1984), in 

tomato (Sinha and McLaren 1989) and in soybeans (Luedders et al. 1978; 

Khan et al. 1986), but these largely remain to be exploited. Transgenic 

cotton lines containing Bacillus thuringiensis toxin genes limited damage to 

initial feeding sites, compared with more extensive skeletonisation in 2 

control cultivars (Flint et al. 1995). Transgenic Bt canola (rape) showed 

excellent resistance to T. ni (Stewart et al. 1996). 

Larvae are easily reared on an artificial diet (e.g. Shorey and Hale 1965; 

Honda et al. 1996). 

The cabbage looper is a major pest of commercial brassicas in North 

America and many other areas where it occurs and causes significant 

damage also, in particular, to lettuce, tomatoes, celery and cotton. Indeed, 

Schwartz (1983) claimed that, if uncontrolled, 92% loss would be sustained 

in the cotton yield in USA, compared with 30% if controlled. Larvae chew 

large irregular holes, leaving only main veins, in the outer leaves of cabbage, 

cauliflower and related plants, often leaving them riddled with holes. Later, 

the outer layers of cabbage heads are eaten and masses of faecal pellets 

contaminate the feeding sites. So much leaf tissue is eaten that heads of 

cabbage and cauliflower are stunted and other leafy vegetables are rendered 

unfit to eat. 

Damage caused to cotton by the larvae consuming leaves is often 

considered less serious, since it frequently occurs late in the growth of the 

cotton plant, so that it may not have a major effect on yield. Cabbage looper 

larvae are essentially foliage feeders and cause their damage in this way. 


Over the last few decades T. ni has become a very widely used laboratory 

insect, particularly in North America. As a result, there are many papers

4.16 


321 

describing laboratory experiments in which parasitoids, predators and/or 

pathogens have been tested on T. ni eggs or larvae. It is often not possible to 

determine from these accounts whether or not the natural enemy involved 

has been found attacking eggs or larvae of T. ni in the field and hence 

possibly a useful control agent. If T. ni is susceptible to the agent in the 

laboratory, such records have often been included, although behavioural or 

other factors might well influence its effectiveness under field conditions. 

No attempt has been made to include reports of all of the minor natural 

enemies, especially in the earlier literature. 

As will become clearer when the situation is discussed later, many 

parasitoids (Table 4.16.1), predators (Table 4.16.2) and pathogens (Table 

4.16.3) have been recorded attacking the cabbage looper in the field and/or 

in the laboratory and there is little doubt that looper populations are 

frequently kept at sub-economic levels by their action. A number of the 

natural enemies recorded are not widespread in distribution and appear to be 

incidental records. 

Many of the predators are widely polyphagous and attack insect pests in 

several orders, although a few are considerably more selective than that. 

Parasitoids, on the other hand, tend to be significantly less polyphagous than 

predators and some appear to be confined to T. ni, 

at least in certain crops. 

Where parasitoids have a relatively broad host range, available records 

indicate that this extends mainly to larvae of other Lepidoptera (generally 

pest species) in the same crop situation. The extent to which it extends also to 

non-pest, non-target species is not documented. Nevertheless, there are 

several species that merit serious consideration as candidates for classical 

biological control. 

All of the Trichogramma species oviposit and develop in the host egg. A 

few others (eg. Copidosoma truncatellum, 

Chelonus blackburni, Chelonus 

insularis) 

oviposit in the egg and develop in the host larva; and the remainder 

are larval and/or pupal parasitoids. 

Naturally occurring epizootics of nuclear polyhedrosis virus in medium 

to large T. ni larvae are considered to be the major mortality factor affecting 

them on cabbage in summer and autumn in southern California (Oatman and 

Platner 1969), on broccoli in Virginia (Hofmaster 1961) and on cabbage in 

North Carolina (Elsey and Rabb 1970b). Polyhedrosis was seldom a major 

factor on cotton in southern California (Ehler 1977b), although outbreaks 

did occur late in the season or at times of high T. ni abundance (Ehler and van 

den Bosch 1974). 

Although the mortality produced is probably not significant, birds and 

bats are known to feed on moths in flight; and earwigs on adults resting 

beneath host plants (McKinney 1944; Sutherland 1966).

Table 4.16.1 

Parasitoids of Trichoplusia ni 

Species 

HYMENOPTERA 

BRACONIDAE 


Cardiochiles nigriceps 

USA Harding 1976 

Chelonus blackburni 

USA Fye & Jackson 1973; Jackson et al. 1979 

Chelonus curvimaculatus 

USA Soldevila & Jones 1991, 1994 

Chelonus nr curvimaculatus 

USA Jones et al. 1981, 1990; Jones 1986 

Chelonus formosanus 

Taiwan Chou 1981 

Chelonus insularis 

USA Ehler et al. 1973; Ehler & van den Bosch 1974; Jones 1982; Henneberry et al. 

1991 

Chelonus sp. USA BŸhler et al. 1985 

Cotesia autographae 

USA Muesebeck & Krombein 1951 

Cotesia congregata 

USA Riley 1883 

Cotesia glomerata 

USA Muesebeck & Krombein 1951; van den Bosch & Hagen 1966; Ehler 1977a 

Cotesia laeviceps 

USA Oatman et al. 1983a 

Cotesia marginiventris 

USA van den Bosch & Hagen 1966; Ehler & van den Bosch 1974, Harding 1976, 

Latheef & Irwin 1983; Henneberry et al. 1991 

Cotesia plutellae 

India Manjunath 1972; Joshi & Sharma 1974 

Cotesia ruficrus 

USA 

India 

McCutcheon et al. 1983 

Manjunath 1972 

Cotesia spp. USA Harding 1976 

Cotesia yakutatensis 

Meteorus autographae 

USA 

India 

Miller & West 1987 


USA Muesebeck & Krombein 1951; Grant & Shepard 1984 


Table 4.16.1 (contÕd) Parasitoids of Trichoplusia ni 

Species 

HYMENOPTERA 

BRACONIDAE (contÕd) 


Meteorus laphygmae 


Microgaster (= Microplitis) 

brassicae USA McKinney 1944; van den Bosch & Hagen 1966; Clancy 1969; Oatman & Platner 

1969; Ehler & van den Bosch 1974; Harding 1976; Ehler 1977a; Jones 1982; 

Oatman et al. 1983a; Henneberry et al. 1991 

Microgaster plutellae 

USA Oatman & Platner 1969; Oatman et al. 1983a 

Microplitis alaskensis 

USA Butler 1958a 

Rogas granulatus 

USA De Gant 1930 

Rogas molestus 

USA Butler 1958a 

Rogas rufocoxalis 

USA McKinney 1944 

Rogas sp. USA Wall & Berberet 1975 

Snellenius manilae 

CHALCIDIDAE 

Taiwan Chou 1981 

Brachymeria intermedia 

Italy 

Dindo 1993 

USA 

Thompson 1980 

Brachymeria lasus 

USA Thompson 1983a,b 

Brachymeria ovata 

USA Elsey & Rabb 1970b; Harding 1976; Patana et al. 1978; Chamberlin & Kok 

1986; Grant & Shepard 1987 

Spilochalcis flavopicta 


Spilochalcis side 


Spilochalcis sp. nr mariae 


4.16 


323


Species 

HYMENOPTERA 


ENCYRTIDAE 

Copidosoma floridanum USA Strand et al. 1991; Baehrecke et al. 1993; Grbk et al. 1992; Ode & Strand 1995 

Copidosoma sp. USA Ehler 1977a 

Copidosoma truncatellum Canada 

USA 

Brazil 

Harcourt 1963 

Riley 1883; McKinney 1944; Pimentel 1961; Oatman 1966; van den Bosch & 

Hagen 1966; Clancy 1969; Oatman & Platner 1969; , Ehler & van den Bosch 

1974; Ehler 1977a; Latheef & Irwin 1983; Oatman et al. 1983a; Roltsch & Mayse 

1983; Chamberlin & Kok 1986 

Silva & Santos 1980 

EULOPHIDAE 

Baryscapus galactopus USA Peck 1963 

Euplectrus comstockii USA McKinney 1944; Harding 1976; Coudron et al. 1994 

Euplectrus platyhypenae USA Wall & Berberet 1974, 1975; Coudron et al. 1990; Kelly & Coudron 1990, 

Euplectrus sp. Santiago van Harten & Miranda 1985 

Pediobius facialis USA Oatman & Platner 1969 

Pediobius nr facialis USA Parkman et al. 1983 

ICHNEUMONIDAE 

Angitia insularis USA Hayslip et al. 1953; Harding 1976 

Campoletis flavicincta USA Krombein et al. 1979; Oatman et al. 1983a 

Campoletis sonorensis USA Cook et al. 1984 

Campoletis sp. USA Oatman et al. 1983a 

Campoletis websteri USA Harding 1976 

Casinaria infesta USA Harding 1976 

Cryptus rutovinctus USA Krombein et al. 1979 



Species 

HYMENOPTERA 


ICHNEUMONIDAE (contÕd) 

Diadegma insulare USA Martin et al. 1982 

Diadegma plutellae USA Harding 1976 

Diadegma spp. USA Sutherland 1966 

Echthromorpha punctum India Manjunath 1972 

Enicospilus sp. India Manjunath 1972 

Gambrus ultimus USA Chamberlin & Kok 1986 

Gelis tenellus USA Sutherland 1966 

Hyposoter exiguae USA Oatman 1966; van den Bosch & Hagen 1966; Clancy 1969; Oatman & Platner 

1969, Ehler & van den Bosch 1974; Ehler 1977a; Jones 1982; Oatman et al. 

1983a, 

Iseropus stercorator orgyiae USA Sutherland 1966 

Itoplectis conquisator USA 

Canada 

Muesebeck & Krombein 1951 

Harcourt 1963 

Microcharops bimaculata Brazil Silva & Santos 1980 

Microcharops tibialis USA Harding 1976 

Nepiera fuscifemora USA Clancy 1969; Oatman et al. 1983a 

Netelia sp. USA Watson et al. 1966 

Patrocloides montanus USA Clancy 1969; Ehler & van den Bosch 1974; Ehler 1977a; Fox et al. 1996 

Pimpla aequalis USA Sutherland 1966 

Pristomerus sp. USA Harding 1976 

Pristomerus spinator USA Harding 1976 

4.16 Trichoplusia ni 325


Species 

HYMENOPTERA 


ICHNEUMONIDAE (contÕd) 

Pterocormus gestuosus USA Mitchell 1961 

Stenichneumon culpator 

cincticornis 

Canada 

USA 

Harcourt 1963 

Chamberlin & Kok 1986 

Vulgichneumon brevicinctor USA Chamberlin & Kok 1986 

PTEROMALIDAE 

Pediobius nr sexdentatus USA Oatman & Platner 1969; Oatman et al. 1983a 

SCELIONIDAE 

Telenomus solitus Guatamala Johnson 1983; Navasero & Oatman 1989 

Telenomus sp. USA Martin et al. 1984 


Trichogramma australicum India Manjunath 1972 

Trichogramma brevicapillum Brazil 

USA 

Hohmann et al. 1988b 

Hoffmann et al. 1990 

Trichogramma chilotraeae India Manjunath 1972 

Trichogramma deion Brazil 

USA 

Hohmann et al. 1988a,b 


Trichogramma evanescens USA Oatman et al. 1968 

Trichogramma exiguum USA Martin et al. 1982; Roltsch & Mayse 1983; Campbell et al. 1991 

Trichogramma japonicum India Manjunath 1972 

Trichogramma minutum USA McKinney 1944; Marston & Ertle 1973; Manweiler 1986 

Trichogramma platneri Brazil 

USA 

Hohmann et al. 1988a,b 

Manweiler 1986 



Species 

HYMENOPTERA 


TRICHOGRAMMATIDAE (contÕd) 

Trichogramma pretiosum USA Oatman 1966; Hoffmann et al. 1975, 1990; Martin et al. 1976b; Ehler 1977a; 

Oatman & Platner 1978; Butler & Lopez 1980; Oatman et al. 1983a 

Trichogramma semifumatum USA Ehler & van den Bosch 1974 

Trichogramma spp. USA van den Bosch & Hagen 1966; Harding 1976; Jones 1982 

Trichogramma thalense Brazil 

USA 

DIPTERA 

Hohmann et al. 1988b 


SARCOPHAGIDAE 

Sacrodexia sternodontis USA Aldrich 1927 

Senotainia sp. USA Manjunath 1972 

TACHINIDAE 

Achaetoneura archippivora USA Butler 1958b 

Aplomya theclarum USA Harding 1976 

Archytas californiae USA van den Bosch & Hagen 1966 

Bessa remota Malaysia Jayanth & Nagarkatti 1984 

Carcelia sp. USA 

India 

Harding 1976 


Chetogena sp. USA van den Bosch & Hagen 1966; Chamberlin & Kok 1986 

Compsilura concinnata Canada 

USA 

Harcourt 1963 

Schaffner & Griswold 1934 

Eucelatoria armigera USA Butler 1958b, Clancy 1969, Henneberry et al. 1991, Oatman 1966, Oatman & 

Platner 1969, van den Bosch & Hagen 1966 

Eucelatoria nr armigera USA Harding 1976, Henneberry et al. 1991 



Species 

DIPTERA 



Eucelatoria rubentis USA Watson et al. 1966 

Euphorocera spp. USA Harding 1976 

Euphorocera tachinomoides USA Oatman 1966; Harding 1976 

Hypantrophaga sp. USA Harding 1976 

Lespesia achaetoneura USA Oatman 1966 

Lespesia archippivora USA Watson et al. 1966; Oatman & Platner 1969; Henneberry et al. 1991 

Lespesia sp. USA Clancy 1969; Chamberlin & Kok 1986 

Madremyia saundersii USA Oatman 1966; Oatman & Platner 1969 

Metachaeta (=Periscepsia) helymus USA Clancy 1969 

Phorocera sp. USA Sutherland 1966 

Phryxe vulgaris USA Sutherland 1966 

Sarcophaga spp. USA van den Bosch & Hagen 1966 

Schizocerophaga leibyi USA Harding 1976 

Siphona plusiae USA Clancy 1969; Harding 1976; Henneberry et al. 1991 

Siphona sp. USA Oatman & Platner 1969; Oatman et al. 1983a 

Voria edentata India Manjunath 1972 

Voria ruralis USA McKinney 1944; Butler 1958b; Pimentel 1961; van den Bosch & Hagen 1966; 

Oatman 1966; Clancy 1969; Oatman & Platner 1969; Elsey & Rabb 1970a; 

Ehler & van den Bosch 1974; Wall & Berberet 1975; Harding 1976; Ehler 1977a; 

Jones 1982; Latheef & Irwin 1983; Oatman et al. 1983a; Chamberlin & Kok 

1986; Gordon et al. 1987; Henneberry et al. 1991; Biever et al. 1992 

Winthemia nr montana USA Harding 1976 

Winthemia quadripustulala USA Allen 1925 



Species 

DIPTERA 



Winthemia rufopicta USA Chamberlin & Kok 1986 

Winthemia spp. USA Elsey & Rabb 1970b 

Zenilla blanda blanda USA West 1925 


Table 4.16.2 Some predators of Trichoplusia ni in USA 

Species Reference 

DERMAPTERA 

LABIDURIDAE 

Labidura riparia Strandberg 1981a,b 

Tawfik et al. 1972 

HEMIPTERA 

ANTHOCORIDAE 

Orius insidiosus Lingren et al. 1978 

Orius tristicolor Ehler et al. 1973; Ehler & van den Bosch 1974; Ehler 1977a; Wilson & Gutierrez 1980; 

Jones 1982; Jones et al. 1983 

LYGAEIDAE 

Geocoris pallens van den Bosch & Hagen 1966; Ehler et al. 1973; Ehler & van den Bosch 1974; Ehler 

1977a; Wilson & Gutierrez 1980 

Geocoris punctipes van den Bosch & Hagen 1966; Barry 1973; Barry et al. 1974; Walker & Turnipseed 

1976; Wilson & Gutierrez 1980; Reed et al. 1984 

NABIDAE 

Nabis alternatus van den Bosch & Hagen 1966; Barry 1973; Barry et al. 1974 

Nabis americoferus van den Bosch & Hagen 1966; Ehler et al. 1973; Ehler & van den Bosch 1974; Ehler 

1977a; Stoltz and Stern 1979; Wilson & Gutierrez 1980 

Nabis roseipennis Reed et al. 1984 

PENTATOMIDAE 

Alcaeorrhynchus grandis McLain 1979 

Euthyrhynchus floridanus McLain 1979 


Table 4.16.2 (contÕd) Some predators of Trichoplusia ni in USA 


Podisus maculiventris Hayslip et al. 1953; Ignoffo et al. 1977; Marston et al. 1978; Richman & Whitcomb 

1978; McLain 1979; Biever et al. 1982 

Stiretrus anchorago Richman 1977 

HEMIPTERA 

REDUVIIDAE 

Sinea complexa van den Bosch & Hagen 1966 

Sinea confusa van den Bosch & Hagen 1966 

Sinea diadema van den Bosch & Hagen 1966 

Sycanus indagator Greene & Shepard 1974 

Zelus bilobus Hayslip et al. 1953 

Zelus exsaguis Whitcomb and Bell 1964 

Zelus renardii van den Bosch & Hagen 1966 

Zelus tetracanthus van den Bosch & Hagen 1966 

NEUROPTERA 

CHRYSOPIDAE 

Chrysopa lanata Ru et al. 1975 

Chrysopa nigricornis van den Bosch & Hagen 1966 

Chrysopa rufilabris Ru et al. 1976 

Chrysopa spp. Pimentel 1961 

Chrysoperla carnea van den Bosch & Hagen 1966; Barry 1973; Barry et al. 1974; Ehler & van den Bosch 

1974; Ehler 1977a; Wilson & Gutierrez 1980 

HEMEROBIIDAE 

Hemerobius spp. van den Bosch & Hagen 1966 




COLEOPTERA 

CARABIDAE 

Calosoma affine van den Bosch & Hagen 1966 

Calosoma peregrinator McKinney 1944 

Labia analis Reed et al. 1984 

COLEOPTERA 

COCCINELLIDAE 

Ceratomegilla maculata fuscilabris Pimentel 1961 

Coccinella novemnotata franciscana van den Bosch & Hagen 1966 

Coccinella transversoguttata Pimentel 1961 

Cycloneda sanguinea van den Bosch & Hagen 1966; Jones 1982 

Hippodamia convergens van den Bosch & Hagen 1966; Jones 1982; Jones et al. 1983 

Hippodamia parenthesis van den Bosch & Hagen 1966 

Hippodamia quinquesignata punctulata van den Bosch & Hagen 1966 

Olla v-nigrum van den Bosch & Hagen 1966 

Paranaemia vittegera van den Bosch & Hagen 1966 

MELYRIDAE 

Collops marginellus van den Bosch & Hagen 1966 

Collops vittatus van den Bosch & Hagen 1966 

DIPTERA 

SYRPHIDAE 

Mesograpta marginata Pimentel 1961 

Sphaerophoria cylindrica Pimentel 1961 

Sphaerophoria menthastri Pimentel 1961 




DIPTERA 

SYRPHIDAE (contÕd) 

Sphaerophoria robusta Pimentel 1961 

Several species van den Bosch & Hagen 1966 

HYMENOPTERA 

VESPIDAE 

Mischocyttarus flavitarsis Bernays & Montelor 1989 

Polistes apachus Cornelius 1993 

Polistes metricus van den Bosch & Hagen 1966; Hunt 1984; Greenstone & Hunt 1993 

Vespula pensylvanica Warren 1990 

SPIDERS 

Misumena ratia Lockley et al. 1989 

Oxyopes salticus Reed et al. 1984; Lockley & Young 1988 

Pardosa spp. Reed et al. 1984 

Phidippus regius Edwards & Jackson 1993, 1994 

Phidippus spp. Edwards & Jackson 1993, 1994 

BIRDS 

Dendroica palmarum Strandberg 1981a 

Passerculus sandwichensis Strandberg 1981a 



More than 20 species of microorganism (viruses, bacteria, protozoa and 

fungi) are associated with T. ni, most of them isolated from field 

populations, but some from laboratory cultures. All were initially isolated 

from larvae, although at least one species of each group has also been 

isolated from pupae, adults and even eggs (Table 4.16.3). Details concerning 

the causative agents and the symptoms they produce are given by Ignoffo 

and Hostetter (1984). Many papers have been published in recent years to 

add details to the records in Table 4.16.3, especially in the field of viruses, 

but also adding to the range of other pathogens, such as a rickettsia-like 

organism (Browning et al. 1982). Naturally-occurring virus infection has 

been found to be a major mortality factor of T. ni larvae in the field, 

particularly late in the season when populations are high. A singleembedded 

nuclear polyhedrosis has been mass produced and applied with 

excellent results by a number of authors to several crops (Ignoffo and 

Hostetter 1984). 

Table 4.16.3 Major pathogens of Trichoplusia ni (Ignoffo and Hostetter 

1984) 

VIRUSES single-embedded nuclear polyhedrosis 

multiple-embedded nuclear polyhedrosis 

granulosis 

cytoplasmic polyhedrosis 

BACTERIA Bacillus thuringiensis 

Serratia marcescens 

PROTOZOA Nosema trichoplusiae 

Thelohania sp. nr. diazoma 

FUNGI Aspergillus flavus 

Beauveria bassiana 

Entomopthora gammae 

Entomopthora sphaerosperma 

Metarrhizium anisopliae 

Metarrhizium brunneum 

Nomuraea rileyi

4.16 Trichoplusia ni 335 

Introductions for biological control of T. ni 

There do not appear to have been any introductions of natural enemies 

specifically for cabbage looper, but rather for the complex of lepidopterous 

larvae with which it is almost always associated. Examples of such 

introductions are shown in Table 4.16.4. 

BRAZIL 

The natural enemies of T. ni larvae on cotton at 3 sites in Paran‡ Province 

included the fungus, Nomuraea rileyi (which killed 76% of larvae at one 

site), a virus disease (that killed up to 47% at two sites), the parasitoids 

Copidosoma truncatellum (reared from about 5% of larvae at 2 sites) and 

Microcharops bimaculata (reared from 7.5% of larvae at 1 site) and the 

fungus Entomopthora sp. (which killed 2.5% of larvae at 1 site) (Silva and 

Santos 1980). The natural enemies of T. ni on cotton in Mato Grosso are 

discussed by Bleicher et al. (1985) and on tomato in Sao Paulo by Gravena 

(1984). 

CARIBBEAN 

T. ni is usually a minor pest of Brassicaceae, although outbreaks 

occasionally cause serious defoliation of crops. A large number of predators 

attack larvae, in addition to the parasitoids that are listed in Table 4.16.5.

Table 4.16.4 Introductions for the biological control of lepidopterous larvae including Trichoplusia ni 

Species 

HYMENOPTERA 

BRACONIDAE 

From To When Result Reference 

Cotesia marginiventris Cape Verde Is 1981 + Lima & van Harten 1985 

Cotesia plutellae India Barbados, Jamaica 1969 + Alam 1992 

Cotesia ruficrus Australia USA 1981 ? McCutcheon et al. 1983 

Microplitis (= Microgaster) demolitor Australia USA 1981 ? Shepard et al. 1983; 

Norlund & Lewis 1985 

Microgaster rufiventris 

ENCYRTIDAE 

Egypt USA 1983 Ð McCutcheon & Harrison 1987 

Copidosoma floridanum 

EULOPHIDAE 

India Barbados pre 1985 + Alam 1992 

Pediobius nr facialis Japan USA pre 1983 * Parkman et al. 1983 

* no indication of field release 



Table 4.16.5 Parasitoids and a fungus attacking T. ni in the Caribbean 

(Alam 1992) 

Species % Parasitisation 

Jamaica Barbados 

BRACONIDAE 

Cotesia sp. (glomerata group) 20.0 

Cotesia plutellae 29.6Ð70.0 3.5 

Glyptapanteles sp. (vitripennis group) 

CHALCIDIDAE 

0.5Ð 2.0 

Brachymeria sp. 2.4 

Brachymeria ovata 

ENCYRTIDAE 

0.5 

Copidosoma sp. 12.5 

Copidosoma floridanum 0.5Ð5.0 

Copidosoma (truncatellum group) 

EULOPHIDAE 

25.8 

Euplectrus platyhypenae 

TACHINIDAE 

Winthemia nr pinguis 

4.2 

and Winthemia nr pyrrhopyga 20.2Ð35.8 

Winthemia sp. 1 specimen only 

ENTOMOPHTHORALES 9.5Ð80.0 

NORTH AMERICA 

The cabbage looper is a widespread and often highly destructive pest of 

cabbage and other Brassicaceae southwards in North America, from about 

the level of Ontario in Canada. Throughout this range it is associated with up 

to about a dozen other species of Lepidoptera. It is third in importance to the 

cabbage white butterfly, Pieris rapae, and the diamondback moth Plutella 

xylostella in Canada (Harcourt 1963) and New York State (Pimentel 1961) 

and about as important as these in southwestern USA (Oatman and Platner 

1969; Reid and Cuthbert 1957). 

In Ontario the encyrtid wasp Copidosoma truncatellum is the most 

important parasitoid and populations are frequently destroyed by a 

polyhedral virus (Harcourt 1963). In New York State a polyhedral virus 

(40% mortality) was the major factor affecting T. ni populations in 1957 but 

less important (7%) in 1958 when predators (especially spiders), caused 2% 

to 3% mortality (Pimentel 1961). In northwestern USA up to 14% of T. ni 

larvae were parasitised on cabbage by the tachinid fly, Voria ruralis (Biever 

et al. 1992). In southern California up to 39% (av. 7.8%) of T. ni eggs were 

parasitised by Trichogramma pretiosum, which was also reared from 

Plutella xylostella eggs. Twelve species of parasitoid were reared from T. ni


larvae and pupae, 7 of which were Hymenoptera and 5 Diptera (Table 

4.16.1). These produced an average of 38.9% mortality, with a maximum of 

66.7% in late autumn. The tachinid, Voria ruralis, was the dominant 

parasitoid, especially during autumn and winter months. The ichneumon, 

Hyposoter exiguae, and the encyrtid, Copidosoma truncatellum, occurred 

most commonly during the summer and autumn months, when the latter was 

associated with a nuclear polyhedrosis virus. Together (and particularly the 

virus) they were responsible for most of the 60% larval mortality. Pupal 

mortality was low (2.0%) and was due to the pteromalid Pediobius 

sexdentatus (Oatman and Platner 1969). Also in southern California, Clancy 

(1969) reared 5 wasps and 5 tachinid flies (Table 4.16.1) from T. ni larvae 

collected from annual weeds (Malva sp., Chenopodium sp.), mustard, tree 

tobacco (Nicotiana glauca) and lucerne. The fly, Voria ruralis, was the most 

abundant parasitoid in autumn and winter and Copidosoma truncatellum the 

most important wasp, the total parasitisation from all species ranging from 

29.5% to 41.1%. A nuclear polyhedrosis virus killed 70.9% of larvae 

collected in summer from weeds and 63.8% from lucerne (Table 4.16.6). 

The mortality caused by the various natural enemies varied according to the 

season and host plant. 

Henneberry et al. (1991) recorded 12 species of parasitoid from larvae of 

the loopers T. ni and Autographa californica on lettuce, lucerne, sugarbeet 

and cotton in southern California. Of these, the braconid Microgaster 

brassicae (30%) Voria ruralis (23%) and Copidosoma truncatellum (23%) 

were the most abundant, with parasitisation rates ranging from 0% to 91.8% 

according to season and crop. Average mortality from viral infection ranged 

between 0.6% and 7.2%, also depending upon the crop. 

Thirteen parasitoid species reared from T. ni on tomatoes in southern 

California (Table 4.16.1) caused mean parasitisation rates of larvae of 51.4% 

and 70.5% and of eggs of 24.6% and 53.4% respectively in two successive 

years. Hyposoter exiguae and Copidosoma truncatellum were the most 

abundant larval parasitoids and Trichogramma pretiosum the most 

important species attacking eggs. The data on population trends and 

percentage parasitisation suggested that there was a density-dependent 

relationship between T. ni and its parasite complex on tomato (Oatman et al. 

1983a). 

In Arizona, the most abundant parasitoid of T. ni larvae collected from 

weeds and cultivated crops was Voria ruralis, which was present throughout 

the year, with peak abundance (up to 100%) in late autumn and winter 

(McKinney 1944; Butler 1958b; Brubaker 1968). Five wasps and 1 tachinid 

fly were reared from larvae, 1 tachinid from pupae and Trichogramma

Table 4.16.6 Natural enemies of Trichoplusia ni larvae in southern California from weed hosts, lucerne and tree tobacco 

(Nicotiana glauca) (from Clancy 1969) 

Month No reared % pupating % killed by 

virus all parasites Voria ruralis all other 

Tachinidae 

Copidosoma 

truncatellum 

all other 

Hymenoptera 

1966 Collections from annual weeds 

May 94 2.1 63.8 34.0 4.3 11.7 10.6 7.4 

June 286 12.6 57.3 30.1 7.7 9.8 4.9 7.0 

July 327 4.6 70.9 24.2 4.0 10.1 5.5 4.6 

Aug 280 6.1 44.3 49.6 16.1 6.4 25.4 1.8 

Sept 193 18.6 31.6 49.7 12.4 9.3 30.0 

Oct 267 41.9 37.8 20.2 6.4 3.4 10.5 

Nov 162 47.5 25.9 26.5 9.3 1.2 16.0 

Dec 

1967 

151 4.6 39.1 56.3 38.4 0.7 16.6 0.7 

Jan 138 31.9 16.7 51.4 34.1 0.7 16.7 

Feb 63 61.9 11.1 27.0 11.1 15.9 

March 7 14.3 28.6 57.1 14.3 42.9 

April 12 41.7 33.3 25.0 25.0 

May 120 60.0 15.8 24.2 16.7 1.7 5.8 

Totals 2100 22.0 42.5 35.2 13.1 5.9 13.3 2.8 

Collections from lucerne 

149 6.7 63.8 29.5 6.0 20.8 0.7 2.0 

Collections from tree tobacco, Nicotiana glauca 

479 47.0 11.9 41.1 21.5 1.0 11.7 6.9 



minutum from eggs collected from lettuce. In addition, the carabid beetle, 

Calosoma peregrinator, fed readily on T. ni larvae (McKinney 1944). 

A study in southern Texas of the loopers T. ni and Chrysodeixis 

includens on a range of host plants revealed high levels (58% to 71%) of 

mortality of larvae and pupae by 29 parasitoid species (Table 4.16.1) during 

all but 4 months of the year, all of which existed at rather low levels (Harding 

1976). 

Turning to cotton in southern California, T. ni is a secondary pest and 

there is good evidence that natural enemies, in particular several predators, 

are mainly responsible for its generally low pest status. The 4 major 

predators there are larvae of the green lacewing, Chrysoperla carnea (which 

consume eggs and larvae) adults and nymphs of the bugs Geocoris pallens 

and Orius tristicolor (which prey upon eggs and small larvae) and adults and 

nymphs of the bug Nabis americoferus (which prey upon larvae of all sizes). 

Spiders, mantids, carabids, vespids and reduviid bugs are among predators 

that are also present in smaller numbers (Ehler and van den Bosch 1974; 

Ehler 1977a). 

Eleven species of parasitoid have been reported from T. ni on cotton, of 

which the following are most important, although their combined effect is 

far less than that of the predators: Trichogramma semifumatum (an egg 

parasitoid), Cotesia marginiventris, Hyposoter exiguae and Microgaster 

brassicae (which attack small larvae and kill hosts when of medium size), 

Copidosoma truncatellum (an egg-larval parasitoid, which emerges from 

large larvae or prepupae), Chelonus texanus (an egg-larval parasitoid which 

kills medium sized hosts), Voria ruralis (which lays eggs on medium sized 

larvae and kills large larvae or prepupae), and Patrocloides montanus (a 

larval-pupal parasitoid which usually oviposits in large larvae). Copidosoma 

truncatellum is polyembryonic and Voria ruralis is often gregarious (Ehler 

and van den Bosch 1974). 

A nuclear polyhedrosis virus was the only pathogen shown to cause T. ni 

mortality, particularly late in the season at peak density of T. ni larvae, when 

levels of 50 to 60% mortality have been observed. 

All of the predators mentioned above are widely polyphagous. The first 4 

parasitoids listed above had a restricted host range and the last 4 were host 

specific in the cotton ecosystem. 

Disappearance of eggs and small larvae, assumed to be due to predation, 

was consistently the major mortality factor. Parasitisation of larvae by any 

parasitoid seldom exceeded 30%, the exception being that by Copidosoma 

truncatellum which often reached 50 to 75%. Detailed life table studies of 

T. ni on cotton were reported by Ehler (1977a).


It was suggested that the temporary nature of the cotton crop, and 

sufficient time each season for only 3 generations of the host, left parasitoids 

insufficient time to build up adequate numbers to thoroughly exploit T. ni 

populations. Also, the low density of T. ni due to the intense activity of 

predators early in each generation, impairs successful search by adult 

parasitoids, particularly those that are host specific (Ehler and van den 

Bosch 1974; Ehler 1977a). 

If the reader is perhaps, somewhat uncertain of what conclusions to draw 

from the extensive data in the foregoing accounts, the studies of Jones et al. 

(1983a), dealing with the impact of parasitoids and predators on T. ni 

populations on celery in California, provide valuable insights. He concludes 

that naturally-occuring entomophagous arthropods do, indeed cause 

irreplaceable mortality of T. ni and that they should be considered a key part 

of any Integrated Pest Management program for the crop. Although 

parasitoids (principally Trichogramma spp., Copidosoma truncatellum and 

Voria ruralis, but also Hyposoter exiguae, Microgaster brassicae, Cotesia 

marginiventris and Chelonus insularis) can explain, for this crop, most 

mortality of eggs and of both small and medium sized larvae, it should not be 

concluded that predators are unimportant. It is possible that the additional 

small amount of mortality due to parasitoids is that required to suppress pest 

density to just below damaging levels. The parasitoids involved have a more 

restricted host range than the 2 most important groups of egg predators in 

celery, namely Coccinellidae (Hippodamia convergens and Cycloneda 

sanguinea) and Anthocoridae (Orius tristicolor) which are both widely 

polyphagous. 

Major parasitoid species 

Laboratory and field studies have been published dealing with many of the 

parasitoids listed in Table 4.16.1. Several of these species that have emerged 

as worthy of serious consideration as biological control agents are dealt with 

below. 

Copidosoma truncatellum Hym.: Encyrtidae 

Females of this small wasp oviposit in T. ni eggs of all ages and 

polyembryonic development occurs after hatching of the host larva. The host 

is later killed in the mature larval or prepupal stage. Either one or two eggs 

are inserted in a host egg during oviposition but, if the latter, generally only 

one parasitoid egg is fertile. Offspring are unisexual although, when 2 fertile 

eggs are laid, both males and females may emerge, an average of 1526 wasps 

per parasitised T. ni (Leiby 1926, 1929).


Young female wasps are better than old at searching for eggs. At 14.8¡C 

and 28.9¡C the period from egg to first adult emergence was 122.9 and 22.4 

days respectively and the duration of a generation was 162.7 and 31.2 days. 

The life span of a female wasp fed on a diet of 20% levulose solution was 

30.3 days at 14.8¡C and 28 days at 35.6¡C. Synchronisation of the parasite to 

T. ni was found to be nearly perfect at 25¡C. At some temperatures the 

parasitoid killed host eggs and at others, larvae before the 5th instar: this 

resulted in the death of the contained parasitoids (Stoner and Weeks 1974, 

1976). 

T. ni larvae parasitised by C. truncatellum consumed 35% more food and 

had a 30% higher maximum weight than unparasitised larvae, which raises a 

concern for at least the short-term effect of biological control (Hunter and 

Stoner 1975). The mortality caused by C. truncatellum appeared to be 

density related (Ehler and van den Bosch 1974). Average parasitisation of 

T. ni eggs in the laboratory was 55.3% (McPherson 1993). However, the 

maximum recorded in cotton fields in southern California was 2.5% (Ehler 

1977a). 

C. truncatellum has been reported from larvae of Noctuidae, 

Geometridae, Cossidae and Coleophoridae (Peck 1963). However, Ehler 

(1977a) points out that it has been reported only from T. ni in Californian 

cotton (van den Bosch and Hagen 1966; Ehler and van den Bosch 1974) and 

that it appears to be specific in this environment. C. truncatellum has a 

Holarctic distribution, but its native home is not clear (Peck 1963). 

Cotesia marginiventris Hym.: Braconidae 

More eggs were laid by this generalist larval parasitoid in 2-day-old T. ni 

larvae than in younger or older larvae. The minimum development period 

from oviposition to adult emergence from the host was 6 days (Boling and 

Pitre 1970). Females were significantly more responsive to host odors after 

brief contact with host larval frass or host-damaged cotton leaves (Turlings 

et al. 1989). C. marginiventris from T. ni and 4 other species of noctuid 

larvae were found to contain a non-occluded, filamentous, baculo-like virus 

(Styer et al. 1987). 

Hyposoter exiguae Hym.: Ichneumonidae 

This solitary endoparasitoid is one of 3 main parasitoids of T. ni on cotton 

(Ehler 1977a) and other crops in California and has a modest ability to 

distinguish unparasitised from parasitised hosts (Beegle and Oatman 1975; 

Browning and Oatman 1984). The female prefers to oviposit in late 1st or 

2nd instar T. ni larvae, although all instars are acceptable. When early instars 

are chosen, the host larvae generally die during the 3rd or 4th instar (Ehler 

1977a). Parasites commencing their development in hosts 1-day-old took


13.85 days for development, whereas those starting in 10-day-old larvae 

required only 7.4 days (Smilowitz and Iwantsch 1975; Jowyk and Smilowitz 

1978). The influence of temperature on development is discussed by 

Browning and Oatman (1981). Weight gain of T. ni larvae is severely 

depressed following parasitisation (Smilowitz and Iwantsch 1973; Iwantsch 

and Smilowitz 1975; Thompson 1982). 

Successful parasitisation of T. ni larvae was correlated with host age, 

ranging from 83% to 88% in 1st, 2nd and early 3rd instars and declining in 

older larvae to 27% in mid 5th instar. Females deposited an average of 2.3 

eggs in 1st instar and 1.3 eggs in 2nd instar larvae, superparasitisation 

declining in later instars (Smilowitz and Iwantsch 1975). 

As many parasitoid eggs were laid in virus-infected host larvae as in 

healthy larvae. Of those females that oviposited in infected hosts, 60% 

transmitted infective doses of virus to 6% of healthy hosts subsequently 

exposed to them. Of female parasitoids that developed in virus-infected hosts, 

90% transmitted infective doses to an average of 21% healthy host larvae 

exposed to them. T. ni larvae parasitised by H. exiguae required twice the 

dosage of virus for infection and the parasitoid completed development before 

the host larvae died (Beegle and Oatman 1974, 1975). Washed H. exiguae 

eggs do not develop to maturity on injection into T. ni larvae unless virus or 

fluid from the parasite oviduct is added (Vinson and Stoltz 1986). Because the 

effects of parasitisation by H. exiguae are observable within 24 hours of 

oviposition and prior to hatching of the parasitoid, it is probable that the 

H. exiguae-associated virus, rather than the developing parasitoid itself, is 

responsible for the metabolic changes produced (Thompson 1986). 

Microgaster brassicae Hym.: Braconidae 

This solitary endoparasitoid is an important mortality factor of T. ni larvae 

on both cabbage and cotton. The female usually oviposits in 1st and 2nd 

instar host larvae and the fully-grown parasite larva leaves through the 

lateral abdominal wall of the medium-sized host larvae to spin a greenish or 

grayish cocoon. The host larva often survives for a few days after parasitoid 

emergence (Ehler 1977a). Duration of parasitoid development from egg to 

adult ranges from 17.7 days at 21.2¡C to 10.7 days at 32.2¡C. Adult 

longevity ranged from 55.5 days at 15.5¡C to 12.9 days at 32.2¡C for males 

and from 76.4 days to 13.7 days for females. Total progeny is largest at 

21.1¡C, averaging 73.2 offspring per female. Rearing methods, mating, 

searching, ovipositional behaviour and interactions with other parasitoid 

species have been described (Browning and Oatman 1984, 1985). 

M. brassicae is native to North America and appears to be specific to 

T. ni in cotton, although it is also known from the alfalfa looper Autographa 

californica on lucerne (Ehler 1977a).


Trichogramma minutum Hym.: Trichogrammatidae 

When attacking eggs of T. ni, females reared from T. ni eggs were more 

fecund than those from Sitotroga cerealella and searched over larger areas 

for hosts (Marston and Ertle 1973). At 27¡C and 50% RH T. minutum 

populations increased more rapidly than those of T. platneri. T. minutum 

does not feed from the host egg nor does it superparasitise eggs even when 

hosts are in short supply (Manweiler 1986). 

Trichogramma platneri Hym.: Trichogrammatidae 

After ovipositing in a host egg, females pierce it again and feed from 

exuding droplets of fluid. T. platneri superparasitised hosts when eggs were 

scarce. At about 27¡C and 50% RH, T. platneri populations increased more 

slowly than those of T. minutum (Manweiler 1986). The maximum number 

of progeny bred from a single T. ni egg was 3 and 73% of male progeny 

emerged from the first eggs exposed to a female. Honey was shown to 

increase parasitoid longevity (Hohmann et al. 1988a, b). 

Trichogramma pretiosum Hym.: Trichogrammatidae 

This species can be reared from egg to adult in vitro (Hoffman et al. 1975). 

Females have a preference for young T. ni eggs, although eggs of all ages are 

accepted (Godin and Boivin 1994). A local Missouri, USA strain of 

T. pretiosum successfully parasitised T. ni eggs in field experiments, large 

host eggs producing more adults than small ones and these adults were more 

fecund and active than those from small eggs (Boldt et al. 1973). A Texan 

strain was effective in the laboratory against T. ni eggs, but not in the field. It 

was able to develop in the same T. ni egg as Trichogramma evanescens if 

eggs of both parasitoids were deposited on the same day (Parker and Pinnell 

1972, 1974). 

In Texas, naturally occurring T. pretiosum assisted in controlling T. ni on 

cotton in field cages (Lingren et al. 1978). In southern California, average 

parasitisation of T. ni eggs ranged from 3 to 47% in tomatoes when releases 

were made at the rate of 200 000 to 318 000 adult wasps per 0.4 ha (Oatman 

and Platner 1978). In Florida 3 releases 3 days apart of T. pretiosum at about 

378 000/acre/release in a 1 acre field cage containing 7 crops resulted in 

substantial parasitisation of T. ni eggs and in suppression of larvae (Martin et 

al. 1976b). Laboratory and field cage studies with T. pretiosum were also 

carried out in California (Ashley et al. 1974) where female parasitoids 

produced from T. ni eggs were larger, more fecund and lived longer than 

those from artificial rearing hosts (Plodia interpunctella and Sitotroga 

cerealella) (Bai et al. 1992).


Voria ruralis Dip.: Tachinidae 

Adults mate soon after eclosion and oviposition commences about 9 days 

later. Eggs laid on the host surface hatch within a minute and young larvae 

penetrate the cuticle and enter a muscle fibre. After about 3 days at 24¡C the 

larva pierces a hole in the dorsal wall of the host abdomen through which it 

inserts its posterior spiracles into the air. After rapid growth of the parasitoid, 

the host dies and the parasitoid larva pupates within the host integument 

(Thompson 1915; Brubaker 1968). When V. ruralis oviposits on 1st instars, 

development is slower and mortality higher than in later instars, except the 

late 5th instar. Development was rarely completed when eggs were laid on 

5th instars, unless they were laid on newly moulted individuals. Females laid 

an average of 310 eggs (Elsey and Rabb 1970a). Development time from egg 

to puparium ranged from 5.4 to 12 days, depending upon the temperature, 

and for the pupa 7 to 8 days at 24¡C. Time from egg to adult varied from 19.4 

days at 20¡C to 10.7 days at 30¡C (Brubaker 1968; Jackson et al. 1969). 

Parasitisation by V. ruralis causes large larvae to eat less than normal (an 

average of 47% reduction (Soo Hoo and Seay 1972). Up to 85% 

parasitisation was observed in field cages, depending upon the numbers of 

mated V. ruralis released, with significant superparasitism at high parasitoid 

densities (Soo Hoo et al. 1974). 

V. ruralis is one of 3 major parasitoids of T. ni on crops in Florida (Martin 

et al. 1982) and cotton in Arizona (Werner and Butler 1979) but was present 

only to the extent of 0 to 0.1% in larvae on lucerne in New Mexico (Gordon 

et al. 1987). In northwestern USA, it was the only parasitoid recovered and 

occurred in 0 to 14% of T. ni larvae (Biever et al. 1992). In Virginia, 

V. ruralis was present in 27% of larvae in 1981 and 17% in 1982 

(Chamberlin and Kok 1986). 

V. ruralis can survive, develop in, and emerge from virus infected larvae. 

However, it does not act as a vector, except occasionally as a mechanical one 

under very restricted conditions (Vail 1981). 

V. ruralis is a widespread parasite and has been recorded as far north as 

Finland and as far south as Trinidad. It has been recorded from a range of 

Lepidoptera. In the United States it is known mainly from larvae of various 

Noctuidae, especially T. ni, but less frequently from the beet armyworm, 

Spodoptera exigua, and other associated species (Jackson et al. 1969; Ehler 

1977a). Ehler and van den Bosch (1974) considered V. ruralis to be host 

specific to T. ni in Californian cotton.


Comment 

It has not been feasible, except for a rather more comprehensive cover of 

parasitoids, to include any but the most relevant of the 2000 or so references 

to T. ni in the literature. Further details can be accessed via the bibliographies 

of Sutherland and Sutherland (1972, 1984) for earlier publications and via 

Commonwealth Agricultural Bureau Abstracts and Lingren and Green 

(1984) for much of the more recent literature. 

Although T. ni is often regarded as a secondary pest of crops in its native 

North America, damaging numbers, nevertheless, occur from time to time, 

particularly when its natural enemies are killed or suppressed by broad 

spectrum insecticides applied for associated primary pests. Predators are 

often claimed to be more important than parasitoids in maintaining T. ni at 

sub-economic levels. 

Regrettably most, if not all, of the major predators involved lack the 

degree of specificity nowadays considered necessary for introduction as 

classical biological control agents. For this reason, further consideration is 

restricted to the potential of parasitoids and viruses. Far more is known from 

southern California than elsewhere of the parasitoid species present and their 

interactions. The following discussion is thus somewhat geographically 

biased and it should be borne in mind that additional species in other regions 

may well have desirable characteristics, especially for their respective 

climatic conditions (see later). 

In addition to the egg parasitoids (e.g. Trichogramma pretiosum and 

T. platneri), there are at least 4 other parasitoids worthy of serious 

consideration (Copidosoma truncatellum, Hyposoter exiguae, Microgaster 

brassicae (Hymenoptera) and Voria ruralis (Tachinidae)). 

If host specificity considerations permit clearance of these species for 

introduction to a new area, a decision must still be taken on which, if not all, 

to establish. It may be useful, therefore, to review (and extend) the 

information presented earlier on their attributes and interactions. 

Copidosoma truncatellum oviposits into the host egg, but hatches in the 

larva and takes about 36 days to develop to adult, so it is present throughout 

the entire larval period of its host. Microgaster brassicae oviposits in 1st and 

2nd instar T. ni larvae and requires about 14 days to develop to adult. The 

mature 3rd instar parasitoid larva emerges from late 3rd or early 4th instar 

hosts after feeding for about 9 days. Hyposoter exiguae commonly oviposits 

into late 1st instar T. ni and emerges from late 3rd or early 4th instar hosts. It 

requires about 16 days from egg to adult at 25¡C, the egg-larval period 

averaging about 10 days. Voria ruralis requires about 13 days from egg to 

adult. It will oviposit on all host instars, but development in most successful


when 2nd or 3rd instars are parasitised. Larval development is then 

completed by the end of the 5th host larval instar (Browning and Oatman 

1984). As a result of overlapping life cycles, two or more of these 4 species 

could inhabit a host larvae simultaneously, unless a species was able to 

discriminate between parasitised and unparasitised hosts. 

As many as 10 adult Voria ruralis may emerge from a single host larva. 

When limited numbers of hosts are present, a female lays more than one egg 

on each larva. Excess eggs may result in premature mortality of the host 

larva and any immature parasitoids already within it. External oviposition 

probably prevents the ovipositing female from receiving sensory 

information about the presence of parasite eggs or larvae already within the 

host; and V. ruralis females will continue to oviposit as long as the host larva 

reacts with any movement. 

Copidosoma truncatellum parasitises host eggs of all ages following 

antennal drumming and ovipositor insertion. Females are apparently able to 

discriminate between unparasitised eggs and those parasitised by other 

C. truncatellum females. Microgaster brassicae females insert their 

ovipositors in all host larvae whether or not parasitised by Copidosoma 

truncatellum, although they are weakly deterred from doing so in larvae 

already parasitised by other M. brassicae females or by Hyposoter exiguae. 

Parasitoid eggs were deposited in 91% of hosts previously unparasitised, 

whereas those already parasitised by M. brassicae or H. exiguae showed 

oviposition levels of 10 and 53% respectively, indicating a response to 

sensory information after insertion of the ovipositor. When M. brassicae 

oviposited in larvae already parasitised by C. truncatellum, the latter 

emerged from 77.5% of the larvae, whereas M. brassicae emerged from only 

12.5%. However, when M. brassicae oviposited in larvae containing the 

slower-developing Hyposoter exiguae, the latter emerged from 16.7% of 

larvae and M. brassicae from 76.7%. Hyposoter exiguae showed little 

discrimination between unparasitised larvae and those parasitised by any of 

the other 3 species, although ovipositor insertion did not result in additional 

eggs in larvae already containing H. exiguae. High levels of parasitisation by 

H. exiguae occurred in host larvae already parasitised by C. truncatellum, 

possibly due to the delayed development of the latter. Nevertheless, only 

C. truncatellum emerged from such larvae. Further details of these and other 

interactions under laboratory conditions are given in the valuable paper by 

Browning and Oatman (1984). It is interesting that the parasitoid complex 

attacking T. ni on cotton in the field shows little change from season to 

season in species composition and relative importance. However, this does 

not enable a simple decision to be made on what impact there would be on 

abundance of T. ni (or on plant damage sustained) if one or more of the


species was omitted from an introduction program. This applies particularly 

to C. truncatellum, with its feature of prolonging the feeding period of 

parasitised larvae. 

A further factor to be taken into consideration is the crop or range of 

crops and the climatic conditions under which T. ni control is particularly 

desired, since the effectiveness of various parasitoid species is affected by 

both sets of factors. 

Relevant to this statement are studies in northwest Florida, which 

reported that Cotesia autographae, Cotesia marginiventris and Meteorus 

autographae caused high T. ni mortality during spring and early summer in a 

mixed cropping system (Martin et al. 1982), and work in southern Florida 

reporting high mortality by Diadegma insulare of T. ni larvae on brassicas. 

Furthermore, Stenichneumon culpator cincticornis and Vulgichneumon 

brevicinctor appear important in New York State (Sutherland 1966), 

Brachymeria ovata in North Carolina (Elsey and Rabb 1970b), and 

Achaetoneura archippivora and Eucelatoria armigera in western United 

States (Martin et al. 1984). 

This suggests that some parasitoid species that are not widely distributed 

can act as significant mortality factors under certain climatic or crop 

conditions. They might well have a very restricted host range, and would be 

available for consideration, if required.

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6 Index of scientific names of insects 

Abacetus optimus Peringuey Col.: Carabidae 90 

abdominale, Dactylosternum 

abdominalis, Aphelinus 

abjectella, Drosica 

abjectum, Praon 

abnormis, Leptomastidea 

abrupta, Platysoma 

absinthii, Aphidius 

absinthii, Praon 

acaenovinae, Aphis 

acalephae, Trioxys 

acantha, Pediobius 

Acanthocoris concoloratus Uhler Hem.: Pentatomidae 233 

Acaulona brasiliana Townsend Dip.: Tachinidae 139 

achaeae, Trichogramma 

Achaetoneura archippivora (Williston) Dip.: Tachinidae 327, 348 

achaetoneura, Lespesia 

Achrysocharis Hym.: Eulophidae 251 

Achrysocharis douglasi, 

see Chrysonotomyia douglasi 251 

Achrysocharoides Hym.: Eulophidae 266, 275, 277 

Achrysophagus Hym.: Encyrtidae 294, 297 

acrobates, Telenomus 

Acroclissoides Hym.: Pteromalidae 201, 228 

Acrosternum Hem.: Pentatomidae 208 

Acrosternum aseadum Rolston Hem.: Pentatomidae 208 

Acrosternum gramineum (Fabricius) Hem.: Pentatomidae 205 

Acrosternum hilare (Say) Hem.: Pentatomidae 205, 208, 230 

Acrosternum marginatum Palisot de Beauvois Hem.: Pentatomidae 208 

Acrosternum pennsylvanicum Palisot de Beauvois Hem.: Pentatomidae 208 

acuminatus, Aenasius 

acuta, Chrysodeixis 

Adialytus salicaphis (Fitch) Hym.: Aphidiidae 37, 66 

Adonia variegata (Goeze) Col.: Coccinellidae 43 

aegyptiacus, Prochiloneurus 

aegyptius, Ischiodon 

477


aegyptius, Xanthogramma 

Aenasius Hym.: Encyrtidae 149 

Aenasius acuminatus Kerrich Hym.: Encyrtidae 147 

Aenasius colombiensis Compere Hym.: Encyrtidae 149 

Aenasius theobromae Kerrich Hym.: Encyrtidae 147 

aequalis, Pimpla 

affine, Calosoma 

affinis, Cryptolaemus 

africanus, Mesochorus 

africanus, Stictopisthus 

africanus, Syrphophagus 

Ageniaspis Hym.: Encyrtidae 123, 125, 265 

Ageniaspis citricola Logvinovskaya Hym.: Encyrtidae 257, 265, 272Ð280, 282, 285 

Agonoscelis rutila (Fabricius) Hem.: Pentatomidae 218 

agraules, Pnigalio 

agrestoria, Echthromorpha 

agriae, Trichogramma 

Agrius convolvuli (Linnaeus) Lep.: Sphingidae 1, 2, 9Ð16 

Agromyza Dip.: Agromyzidae 236 

Agromyza obtusa, 

see Melanagromyza obtusa 236 

Agromyza phaseoli, 

see Ophiomyia phaseoli 236 

agromyzae, Neodimmockia 

agromyzae, Sphegigaster 

agromyzae, Trigonogastra 

alaskensis, Microplitis 

albiceps, Phytomyza 

albicollis, Hyperaspis 

albipes, Pleurotropitiella 

albizonalis, Autocharis 

albizonalis, Deanolis 

albizonalis, Noorda 

Alcaeorrhynchus grandis (Dallas) Col.: Pentatomidae 330 

Aleiodes Hym.: Braconidae 21, 23, 28 

Aleiodes aligharensi (Quadri) Hym.: Braconidae 21, 23, 28 

aliberti, Brachymeria 

aligarhensis, Diaphorencyrtus 

aligharensi, Aleiodes 

Allograpta nasuta (Macquart) Dip.: Syrphidae 43

Allograpta pfeifferi, 

see Allograpta nasuta 

Allotropa Hym.: Platygasteridae 150, 301 

Scientific Index 479 

Allotropa citri Muesebeck Hym.: Platygasteridae 295, 298Ð301, 307, 315 

Allotropa kamburovi Annecke and Prinsloo Hym.: Platygasteridae 295, 298 

Allotropa mecrida (Walker) Hym.: Platygasteridae 295, 298, 306 

Alloxysta Hym.: Charipidae 65, 67, 68, 72 

Alloxysta brevis (Thompson) Hym.: Charipidae 72 

43 

Alloxysta darci (Girault) Hym.: Charipidae 72 

aloysiisabaudiae, Trissolcus 

alternatus, Nabis 

alticeps, Leucopis 

amabilis, Eublemma 

Amatellon Hym.: Eulophidae 266, 273, 282 

ambiguus, Lipolexis 

ambiguus, Lysiphlebus 

Amblypelta cocophaga China Hem.: Coreidae 226 

Amblyteles fuscipennis Wesmael Hym.: Ichneumonidae 12 

americana, Asaphes 

americana, Carcinophora 

americana, Psalis 

americoferus, Nabis 

Amobia Dip.: Sarcophagidae 190 

amygdali, Hyalopterus 

amyoti, Glaucias 

Anacanthocoris concoloratus (Uhler) Hem.: Coreidae 224 

Anagyrus Hym.: Encyrtidae 147, 152 

Anagyrus ananatis Gahan Hym.: Encyrtidae 145, 147, 149, 152, 154Ð156 

Anagyrus bohemani (Westwood) Hym.: Encyrtidae 294, 297, 305 

Anagyrus coccidivorus, 

see Anagyrus ananatis 149 

Anagyrus greeni Howard Hym.: Encyrtidae 294, 297 

Anagyrus kivuensis Compere Hym.: Encyrtidae 149, 307 

Anagyrus pseudococci (Girault) Hym.: Encyrtidae 287, 297, 299Ð301, 303, 305Ð307, 

311, 315, 316 

Anagyrus sawadai Ishii Hym.: Encyrtidae 294, 297, 305, 307 

analis, Lebia 

ananatis, Anagyrus 

Anasa tristis (De Geer) Hem.: Pentatomidae 230 

Anastatus Hym: Eupelmidae 203, 211, 214, 215, 223, 227


Anastatus bifasciatus Boyer de Fonscolombe Hym.: Eupelmidae 203, 223 

Anastatus dasyni Ferri re Hym: Eupelmidae 202 

Anastatus japonicus Ashmead Hym: Eupelmidae 203, 225 

anchorago, Stiretrus 

angelicae, Trioxys 

angelicus, Pseudaphycus 

Angita insularis Hym.: Ichneumonidae 324 

Angitia plutellae, 

see Diadegma insulare 

angolensis, Chilocorus 

angustifrons, Pseudaphycus 

Anisochrysa basalis, 

see Mallada basalis 291 

Anisolabis annulipes, see Euborellia annulipes 90 

Ankylopteryx octopunctata Fabricius Neu.: Chrysopidae 271 

annulipes, Anisolabis 

annulipes, Euborellia 

Anochaetus Hym.: Formicidae 92 

anomidis, Apanteles 

Anomis erosa (HŸbner) Lep.: Noctuidae 18 

Anomis flava (Fabricius) Lep.: Noctuidae 1, 2, 17Ð31 

Anoplolepis custodiens (F. Smith) Hym.: Formicidae 305 

Anoplolepis longipes (Jerdon) Hym.: Formicidae 155 

antennata, Nezara 

antennata, Pauesia 

antestiae, Gryon 

antestiae, Hadronotus 

Antestia orbana Kirk Hem.: Pentatomidae 225 

Anthocoris Hem.: Anthocoridae 43 

Antilochus coquebertii (Fabricius) Hem.: Pyrrhocoridae 137, 138 

antinorii, Bogosia 

Antonina graminis (Maskell) Hem.: Pseudococcidae 155 

apachus, Polistes 

Apanteles Hym.: Braconidae 12, 14, 23, 29, 90, 192 

Apanteles anomidis Watanabe Hym.: Braconidae 21, 23, 29, 31 

Apanteles ruficrus, 

see Cotesia ruficrus 23 

Apanteles syleptae Ferri re Hym.: Braconidae 23, 28 

apetzi, Scymnus 

Aphanogmus dictynna (Waterston) Hym.: Ceraphronidae 166, 168, 182 

Aphelinus Hym.: Aphelinidae 37, 51, 63, 64, 68, 72, 82, 83


Aphelinus abdominalis (Dalmer) Hym.: Aphelinidae 37, 50, 61, 65, 67, 68, 71 

Aphelinus asychis Walker Hym.: Aphelinidae 50 

Aphelinus basalis, see Aphelinus abdominalis 

Aphelinus brevis, see Alloxysta brevis 

37, 67 

Aphelinus chaoniae Walker Hym.: Aphelinidae 50 

Aphelinus flavipes, see Aphelinus abdominalis 37, 50, 65, 71 

Aphelinus gossypii Timberlake Hym.: Aphelinidae 37, 50, 68, 71, 72, 83 

Aphelinus humilis Mercet Hym.: Aphelinidae 50 

Aphelinus kashmiriensis, see Aphelinus gossypii 50 

Aphelinus mali (Haldeman) Hym.: Braconidae 50, 63, 68, 72 

Aphelinus mariscusae (Risbec) Hym.: Aphelinidae 37 

Aphelinus nigritus, see Aphelinus varipes 50 

Aphelinus paramali Zehavi and Rosen Hym.: Aphelinidae 50 

Aphelinus semiflavus Howard Hym.: Aphelinidae 56, 69, 72 

Aphelinus varipes (Foerster) Hym.: Aphelinidae 50, 61 

Aphidencyrtus Hym.: Encyrtidae 57, 63, 65 

Aphidencyrtus aligarhensis, see Diaphorencyrtus aligarhensis 121, 131 

Aphidencyrtus aphidiphagus, see Syrphophagus aphidivora 72 

Aphidencyrtus aphidivora, see Syrphophagus aphidivora 42 

Aphidencyrtus diaphorinae, see Diaphorencyrtus aligarhensis 

aphidimyza, Aphidoletes 

aphidiphagus, Aphidencyrtus 

aphidis, Pachyneuron 

121 

Aphidius Hym.: Aphidiidae 38, 53 

Aphidius absinthii Marshall Hym.: Aphidiidae 37, 67 

Aphidius avenae, see Aphidius picipes 63 

Aphidius cardui, see Lysiphlebus fabarum 39 

Aphidius colemani Viereck Hym.: Aphidiidae 37, 51, 60, 61, 64, 66, 68, 73, 81Ð83 

Aphidius ervi Haliday Hym.: Aphidiidae 37, 51 

Aphidius flavipes, see Aphelinus abdominalis 37 

Aphidius floridaensis Smith Hym.: Aphidiidae 51 

Aphidius funebris Mackauer Hym.: Aphidiidae 37 

Aphidius gifuensis Ashmead Hym.: Aphidiidae 51, 68, 74, 83 

Aphidius lonicerae, see Aphidius urticae 51 

Aphidius matricariae (Haliday) Hym.: Aphidiidae 5, 37, 51, 67, 74, 82 

Aphidius phorodontis, see Aphidius matricariae 51, 74 

Aphidius picipes (Nees) Hym.: Aphelinidae 51, 63 

Aphidius platensis, see Aphidius colemani 37, 73


Aphidius ribis Haliday Hym.: Aphidiidae 37 

Aphidius rosae Haliday Hym.: Aphidiidae 38 

Aphidius salicis Haliday Hym.: Aphidiidae 38 

Aphidius similis Starù and Carver Hym.: Aphidiidae 38, 51 

Aphidius sonchi Marshall Hym.: Aphidiidae 51 

Aphidius transcaspicus, see Aphidius colemani 73 

Aphidius urticae Haliday Hym.: Aphidiidae 51 

Aphidius uzbekistanicus Luzhetzki Hym.: Aphidiidae 

aphidivora, Aphidencyrtus 

aphidivora, Syrphophagus 

38, 51 

Aphidoletes aphidimyza (Rondani) Dip.: Cecidomyiidae 49, 57, 69, 79, 80 

Aphis Hem.: Aphididae 75, 76, 78, 79, 82 

Aphis acaenovinae Eastop Hem.: Aphididae 76 

Aphis citricola, see Aphis spiraecola 70, 78 

Aphis craccivora Koch Hem.: Aphididae 1, 2, 33Ð83 

Aphis fabae Scopoli Hem.: Aphididae 73, 75 

Aphis gossypii Glover Hem.: Aphididae 1, 2, 33Ð83 

Aphis nerii Boyer de Fonscolombe Hem.: Aphididae 60, 70, 73, 74, 76Ð78 

Aphis punicae Passerini Hem.: Aphididae 73 

Aphis spiraecola Patch Hem.: Aphididae 66, 70, 76, 78 

Aphis zizyphi Theobald Hem.: Aphididae 

apicalia, Exorista 

apiciflavus, Scymnus 

73 

Apleurotropis Hym.: Eulophidae 266, 273 

Aplomya theclarum (Scudder) Dip.: Tachinidae 327 

Aprostocetus Hym.: Eulophidae 

archippivora, Achaetoneura 

archippivora, Lespesia 

241, 266 

Archytas californiae (Walker) Dip.: Tachinidae 

arcuata, Coccinella 

arcuata, Ectophasiopsis 

argenteopilosus, Eriborus 

327 

Argyrophylax atropivora, see Zygobothria atropivora 

armigera, Eucelatoria 

armigera, Helicoverpa 

12 

Asaphes americana, see Asaphes lucens 72 

Asaphes lucens (Provancher) Hym.: Pteromalidae 72 

Asaphoideus niger Girault Hym.: Pteromalidae 271, 272

Ascotolinx funeralis Girault Hym.: Eulophidae 266, 272, 276 

aseadum, Acrosternum 

asiaticus, Trioxys 

Asolcus mitsukurii, see Trissolcus mitsukurii 224 

asychis, Aphelinus 

atomella, Melanagromyza 

atricornis, Phytomyza 

atropivora, Argyrophylax 

atropivora, Sturmia 

atropivora, Zygobothria 

attrisium, Rhychium 

auberti, Diaphorina 

auctus, Trioxys 

aurantii, Toxoptera 

auratocauda, Cadurcia 

auratocauda, Sturmia 

auropunctata, Wasmannia 

australicum, Trichogramma 

australiensis, Eupelmus 

Autocharis albizonalis, see Deanolis albizonalis 106 

Autographa brassicae, see Trichoplusia ni 318 

Autographa californica (Speyer) Lep.: Noctuidae 338, 342 

autographae, Cotesia 

autographae, Meteorus 

avenae, Aphidius 

Axiagastus campbelli Distant Hem.: Pentatomidae 226 

axyridis, Harmonia 

ayyari, Tetrastichus 

Azteca Hym.: Formicidae 176 

Bactrocera ferrugineus (Fabricius) Dip.: Tephritidae 109 

Bactrocera frauenfeldi (Schiner) Dip.: Tephritidae 109 

balteatus, Episyrphus 

barberi, Sympherobius 

Baryscapus galactopus (Ratzeberg) Hym.: Eulophidae 324 

basalis, Anisochrysa 

basalis, Aphelinus 

basalis, Chrysopa 

Scientific Index 483


basalis, Mallada 

basalis, Trissolcus 

basicurvus, Trioxys 

bella, Leucopis 

Belonuchus ferrugatus (Erichson) Col.: Staphylinidae 92, 93, 102 

Belonuchus quadratus Kraatz. Col.: Staphylinidae 92, 102 

beneficia, Polycystomyia 

Bessa remota (Aldrich) Dip.: Tachinidae 327 

bicarinativentris, Plutarchia 

bicarinatum, Tetramorium 

bicolor, Charops 

bicolor, Galerita 

bicolor, Microdon 

bicolor, Paragus 

bicolor, Propagalerita 

bifasciatus, Anastatus 

biguttatus, Scymnus 

bilobus, Zelus 

bilucenarius, Nephus 

bilucenarius, Scymnus 

bimaculata, Microcharops 

bimaculata, Sturmia 

binaevatus, Scymnus 

Biosteres Hym.: Braconidae 251 

biplaga, Earias 

bipunctatus, Nephus 

bipunctatus, Scymnus 

bipustulatus, Chilocorus 

blackburni, Chelonus 

blanda blanda, Zenilla 

Blepyrus insularis (Cameron) Hym.: Encyrtidae 294, 297 

Blepyrus saccharicola Gahan Hym.: Encyrtidae 294, 297, 300, 307 

Bogosia antinorii Rondani Dip.: Tachinidae 201, 202, 211, 217, 232Ð234 

Bogosia helva (Wiedermann) Dip.: Tachinidae 139 

bohemani, Anagyrus 

bolivari, Neoprochiloneurus 

boninensis, Chrysopa 

boninensis, Mallada


borbonicus, Paragus 

borellii, Labia 

Brachycantha Col.: Coccinellidae 146, 152 

Brachycaudus Hem.: Aphididae 75 

Brachycaudus cardui (Linnaeus) Hem.: Aphididae 75 

Brachymeria Hym.: Chalcididae 21, 24, 27, 191, 192, 337 

Brachymeria aliberti (Schmitz) Hym.: Chalcididae 24, 28 

Brachymeria intermedia (Nees) Hym.: Chalcididae 323 

Brachymeria lasus (Walker) Hym.: Chalcididae 24, 30, 191, 192, 323 

Brachymeria madecassa Steffan Hym.: Chalcididae 24, 29 

Brachymeria multicolor (Kieffer) Hym.: Chalcididae 21, 24, 29 

Brachymeria obscurata, see Brachymeria lasus 24, 192 

Brachymeria ovata (Say) Hym.: Chalcididae 323, 337, 348 

Brachymeria paolii Masi Hym.: Chalcididae 24, 29 

Brachymeria tibialis Steffan Hym.: Chalcididae 21, 24 

Bracon Hym.: Braconidae 190, 192, 265, 276, 282 

Bracon greeni Ashmead Hym.: Braconidae 190, 194 

Bracon phyllocnistidis (Muesebeck) Hym.: Braconidae 265, 274, 282 

brasiliana, Acaulona 

brassicae, Autographa 

brassicae, Brevicoryne 

brassicae, Liriomyza 

brassicae, Microgaster 

brassicae, Microplitis 

brevicapillum, Trichogramma 

brevicinctor, Vulgichneumon 

Brevicoryne brassicae (Linnaeus) Hem.: Aphididae 48 

brevipes, Dysmicoccus 

brevipes, Pseudococcus 

brevipetiolatus, Zaommomentedon 

brevipetioletus, Visnuella 

brevis, Alloxysta 

brevis, Aphelinus 

Brinckochrysa scelestes (Banks) Neu.: Chrysopidae 291 

brochymenae, Trissolcus 

bromeliae, Diaspis 

Brumoides lineatus (Weise) Col.: Coccinellidae 291 

Brumus suturalis (Fabricius) Col.: Coccinellidae 43, 291, 309


brunneicornis, Sphegigaster 

brunnipennis, Plautia 

c-nigrum, Hyperaspis 

cadaverica, Scleroderma 

Cadurcia auratocauda (Curran) Dip.: Tachinidae 

caenosus, Dictyotis 

californiae, Archytas 

californica, Autographa 

californica, Coccinella 

californicus, Ooencyrtus 

22 

Calliceras dictynna, see Aphanogmus dictynna 168 

Calliephaltes Hym.: Ichneumonidae 195 

Callitula filicornis Del. Hym.: Pteromalidae 243 

Callitula viridicoxa (Girault) Hym.: Eurytomidae 243, 251 

Callitula yasudi Yasuda Hym.: Pteromalidae 243 

Calosoma affine Chaudoir Col.: Carabidae 332 

Calosoma peregrinator GuŽrin Col.: Carabidae 332, 340 

Calosoma schayeri Erichson Col.: Carabidae 

campaniformis, Eumenes 

campbelli, Axiagastus 

26 

Camplyocheta Dip.: Tachinidae 22 

Campoletis Hym.: Ichneumonidae 324 

Campoletis flavicincta Ashmead Hym.: Ichneumonidae 324 

Campoletis sonorensis (Cameron) Hym.: Ichneumonidae 324 

Campoletis websteri, see Campoletis sonorensis 324 

Camponotus Hym.: Formicidae 153 

Camponotus friedae Forel Hym.: Formicidae 155 

Campyloneura Hym.: Braconidae 190, 192 

Cantheconidia furcellata, see Eucanthecona furcellata 

capensis, Condica 

capensis, Prospalta 

capicola, Platynaspis 

22 

Carcelia Dip.: Tachinidae 111, 327 

Carcelia cosmophilae (Curran) Dip.: Tachinidae 22, 27 

Carcelia illota (Curran) Dip.: Tachinidae 22, 27 

Carcelia kockiana Tours. Dip.: Tachinidae 22 

Carcinophora americana (Palisot de Beauvois) Derm.: Labiidae 90, 102


cardiae, Diaphorina 

Cardiochiles nigriceps Viereck Hym.: Braconidae 

cardui, Aphidius 

cardui, Brachycaudus 

cardui, Lysiphlebus 

carnea, Chrysopa 

carnea, Chrysoperla 

322 

Carpocoris mediterraneus Tamanini Hem.: Pentatomidae 224 

Carpophilus Col.: Nitidulidae 144 

Casinaria infesta (Cresson) Hym.: Ichneumonidae 324 

Cathartus Col.: Silvanidae 

centaureae, Trioxys 

centrosematis, Ophiomyia 

92, 93, 102 

Cephalonomia Hym.: Bethylidae 175 

Cephalonomia stephanoderis Betrem Hym.: Bethylidae 157, 166, 168, 172Ð177, 182 

Ceraphron dictynna, see Aphanogmus dictynna 

cerasicola, Ephedrus 

182 

Ceratomegilla maculata fuscilabris De Geer Col.: Coccinellidae 

cerealella, Sitotroga 

332 

Cermatulus nasalis (Westwood) Hem.: Pentatomidae 

ceroplastae, Coccophagus 

chaoniae, Aphelinus 

22, 27, 218, 225 

Charichirus Col.: Staphylinidae 92 

Charips, see Alloxysta 65 

Charops Hym.: Ichneumonidae 25 

Charops bicolor (SzŽpligeti) Hym.: Ichneumonidae 12, 14, 21, 25, 29 

Chartocerus walkeri Hayat Hym.: Signiphoridae 120, 123, 125 

Cheilomenes lunata (Fabricius) Col.: Coccinellidae 43 

Cheilomenes sexmaculata Fabricius Col.: Coccinellidae 43, 65, 67, 122, 128 

Cheilomenes sulphurea (Oliver) Col.: Coccinellidae 43 

Cheilomenes vicina (Mulsant) Col.: Coccinellidae 43 

Cheiloneurus Hym.: Encyrtidae 123, 125, 129 

Chelonus Hym.: Braconidae 190, 192, 195, 322 

Chelonus blackburni Cameron Hym.: Braconidae 321, 322 

Chelonus curvimaculatus Cameron Hym.: Braconidae 322 

Chelonus formosanus Sonan Hym.: Braconidae 322 

Chelonus insularis (Cresson) Hym.: Braconidae 321, 322, 341 

Chelonus, texanus Cresson Hym.: Braconidae 340


Chetogena Dip.: Tachinidae 327 

Chilo simplex, see Chilo suppressalis 15 

Chilo suppressalis Walker Lep.: Pyralidae 15 

Chilocorus angolensis Crotch Col.: Coccinellidae 309 

Chilocorus bipustulatus (Linnaeus) Col.: Coccinellidae 

chilonis, Trichogramma 

chilotraeae, Trichogramma 

292, 303 

Chlorocytus Hym.: Pteromalidae 242, 246 

Chloropulvinaria psidii (Maskell) Hem.: Coccidae 

chloropus, Telenomus 

314 

Chromatomyia horticola (Goureau) Dip.: Agromyzidae 239, 252 

Chrysocharis Hym.: Eulophidae 266, 273, 274, 276 

Chrysocharis pentheus (Walker) Hym.: Eulophidae 266, 274, 275 

Chrysodeixis acuta (Walker) Lep.: Noctuidae 28 

Chrysodeixis includens Walker Lep.: Noctuidae 340 

Chrysonotomyia Hym.: Eulophidae 241, 266, 273, 277 

Chrysonotomyia douglasi (Girault) Hym.: Eulophidae 241, 251 

Chrysonotomyia erythraea (Silvestri) Hym.: Eulophidae 241, 251 

Chrysonotomyia formosa (Westwood) Hym.: Eulophidae 241, 251 

Chrysopa Neu.: Chrysopidae 22, 146, 291, 331 

Chrysopa basalis, see Mallada basalis 271 

Chrysopa bipunctatus, see Nephus bipunctatus 

Chrysopa boninensis Okamoto Neu.: Chrysopidae 

Chrysopa carnea, see Chrysoperla carnea 

122, 271, 273, 282, 283 

Chrysopa formosa Brauer Neu.: Chrysopidae 62 

Chrysopa intima MacLachlan Neu.: Chrysopidae 62 

Chrysopa irregularis Banks Neu.: Chrysopidae 146, 152 

Chrysopa kulingensis Navas Neu.: Chrysopidae 190, 192 

Chrysopa lacciperda, see Odontochrysa lacciperda 291, 302, 314 

Chrysopa lanata Banks Neu.: Chrysopidae 331 

Chrysopa nigricornis Burmeister Neu.: Chrysopidae 331 

Chrysopa orestes Banks Neu.: Chrysopidae 65 

Chrysopa pallens Tieder Neu.: Chrysopidae 62 

Chrysopa perla (Linnaeus) Neu.: Chrysopidae 69 

Chrysopa ramburi Schneider Neu.: Chrysopidae 146, 152 

Chrysopa rufilabris Burmeister Neu.: Chrysopidae 331 

Chrysopa scelestes, see Brinckochrysa scelestes 291 

Chrysopa septempunctata, see Chrysopa pallens 62


Chrysopa sinica (Tieder) Neu.: Chrysopidae 62, 69, 271 

Chrysoperla carnea (Stephens) Neu.: Chrysopidae 43, 69, 291, 302, 303, 331, 340 

Chrysoperla sinica, see Chrysopa sinica 62 

Chrysopilus Dip.: Rhagionidae 92 

Chrysopilus ferruginosus (Wiedermann) Dip.: Rhagionidae 92, 93, 95, 96 

Chrysoplatycerus splendens (Howard) Hym.: Encyrtidae 294, 297, 305, 316 

ciliata, Zygobothria 

cinctipennis, Closterocerus 

cingulatus, Dysdercus 

circulus, Halticoptera 

Cirrospiloideus phyllocnistoides, see Citrostichus phyllocnistoides 268 

Cirrospilus Hym.: Eulophidae 102, 241, 266, 272, 273, 275, 278 

Cirrospilus ingenuus Gahan Hym.: Eulophidae 264, 266, 274, 275, 277, 283 

Cirrospilus longefasciatus Ferri re Hym.: Eulophidae 266 

Cirrospilus lyncus Walker Hym.: Eulophidae 266, 274 

Cirrospilus phyllocnistis Ishii Hym.: Eulophidae 266, 277 

Cirrospilus phyllocnistoides, see Citrostichus phyllocnistoides 268 

Cirrospilus pictus (Nees) Hym.: Eulophidae 266, 275, 276 

Cirrospilus quadristriatus (Subba Rao & Ramamani) Hym.: Eulophidae 257, 266, 272, 

273, 275, 277Ð280, 283, 285 

Cirrospilus variegatus (Masi) Hym.: Eulophidae 266, 277 

Cirrospilus vittatus (Walker) Hym.: Eulophidae 266, 275 

cirsii, Trioxys 

citrella, Phyllocnistis 

citri, Allotropa 

citri, Diaphorina 

citri, Kratoysma 

citri, Planococcus 

citricola, Ageniaspis 

citricola, Aphis 

citricola, Psylla 

citriculus, Pseudococcus 

citrisuga, Psylla 

citroimpura, Trioza 

Citrostichus phyllocnistoides (Narayanan) Hym.: Eulophidae 268, 272, 273, 275, 277, 

279, 280, 283, 285 

Clausenia josefi Rosen Hym.: Encyrtidae 294, 297 

Clausenia purpurea Ishii Hym.: Encyrtidae 303


clavata, Gymnosoma 

Cleodiplosis koebelei (Felt) Dip.: Cecidomyiidae 151, 154 

Cleothera Col.: Coccinellidae 150 

Closterocerus Hym.: Eulophidae 177, 273 

Closterocerus cinctipennis Ashmead Hym.: Eulophidae 268, 278 

Closterocerus trifasciatus Westwood Hym.: Eulophidae 268, 275, 277 

Coccodiplosis smithi (Felt) Dip.: Cecidomyiidae 

coccidivora, Triommata 

coccidivorus, Anagyrus 

293 

Coccidoxenoides Hym.: Encyrtidae 299, 301 

Coccidoxenoides peregrinus Timberlake Hym.: Encyrtidae 

304, 305, 307, 311, 312, 315, 316 

287, 294, 296, 297, 300Ð302, 

Coccinella arcuata, see Harmonia octomaculata 60, 300 

Coccinella californica Mann. Col.: Coccinellidae 292 

Coccinella novemnotata francisciana Casey Col.: Coccinellidae 332 

Coccinella repanda, see Coccinella transversalis 60, 292 

Coccinella septempunctata (Linnaeus) Col.: Coccinellidae 26, 43, 62, 65Ð67, 69, 306 

Coccinella semipunctata Col.: Coccinellidae 292 

Coccinella transversalis (Thunberg) Col.: Coccinellidae 60, 292 

Coccinella transversoguttata Faldeman Col.: Coccinellidae 332 

Coccinella undecimpunctata Linnaeus Col.: Coccinellidae 

coccophaga, Amblypelta 

69 

Coccophagus Hym.: Aphelinidae 123, 126 

Coccophagus ceroplastae (Howard) Hym.: Aphelinidae 123, 126 

Coccophagus heteropneusticus Compere Hym.: Aphelinidae 311 

Coccus pseudomagnoliarum (Kuwana) Hem.: Coccidae 

coffea, Phymastichus 

coffeicola, Heterospilus 

cognatoria, Hadrojoppa 

colemani, Aphidius 

299 

Collops marginellus Le Conte Col.: Melyridae 332 

Collops vittatus (Say) Col.: Melyridae 

colombiensis, Aenasius 

communis, Trioxys 

comperei, Telenomus 

complanatus, Trioxys 

complexa, Sinea 

332 

Compsilura concinnata (Meigen) Dip.: Tachinidae 327


comstocki, Pseudococcus 

comstockii, Euplectrus 

concinnata, Compsilura 

concolor, Pachyneuron 

concoloratus, Anacanthocoris 

Condica capensis GuenŽe Lep.: Noctuidae 194 

confrater, Eupeodes 

confrater, Syrphus 

confusa, Sinea 

confusum, Trichogramma 

confusus, Lysiphlebus 

congregata, Cotesia 

Conocephalus saltator (Saussure) Ort.: Tettigoniidae 146 

conquisator, Itoplectis 

constrictus, Scymnus 

convergens, Hippodamia 

convolvuli, Agrius 

convolvuli, Herse 

Copidosoma Hym.: Encyrtidae 195, 324, 337 

Copidosoma floridanum (Ashmead) Hym.: Encyrtidae 324, 326, 337 

Copidosoma truncatellum Dalman Hym.: Encyrtidae 317, 321, 324, 335, 337Ð342, 

346Ð348 

coquebertii, Antilochus 

corbetti, Elasmus 

Coruna Hym.: Pteromalidae 218 

Cosmophila, see Anomis 

Cosmophila erosa, see Anomis erosa 18 

Cosmophila flava, see Anomis flava 18 

Cosmophila indica, see Anomis flava 18 

cosmophilae, Carcelia 

cosmophilae, Zenillia 

Cosmopolites Col.: Curculionidae 86 

Cosmopolites pruinosus Heller Col.: Curculionidae 86 

Cosmopolites sordidus (Germar) Col.: Curculionidae 1, 2, 85Ð104 

Cotesia Hym.: Braconidae 322, 336 

Cotesia autographae Muesebeck Hym.: Braconidae 322, 348 

Cotesia congregata (Say) Hym.: Braconidae 322 

Cotesia glomerata (Linnaeus) Hym.: Braconidae 322, 337


Cotesia laeviceps (Ashmead) Hym.: Braconidae 322 

Cotesia marginiventris (Cresson) Hym.: Braconidae 322, 336, 340Ð342, 348 

Cotesia plutellae Kurdjimov Hym.: Braconidae 322, 336, 337 

Cotesia ruficrus (Haliday) Hym.: Braconidae 23, 322, 336 

Cotesia yakutatensis Ashmead Hym.: Braconidae 

craccivora, Aphis 

crassipennis, Ectophasia 

crawfordi, Ophelosia 

322 

Cremastus flavoorbitalis, see Trathala flavoorbitalis 193 

Cremastus hapaliae Cushman Hym.: Ichneumonidae 191 

Crematogaster Hym.: Formicidae 153 

Crematogaster curvispinosa Mayr Hym.: Formicidae 

cristatus, Telenomus 

169, 175 

Cristicaudus nepalensis (Takada) Hym.: Aphidiidae 

crossota, Plautia 

crypticus, Trissolcus 

53 

Cryptochetum sp. Dip.: Cryptochetidae 293 

Cryptogonus orbiculus, see Nephus bipunctatus 296 

Cryptolaemus Col.: Coccinellidae 146, 150, 152, 155 

Cryptolaemus affinis Crotch Col.: Coccinellidae 153, 292, 304 

Cryptolaemus montrouzieri Mulsant Col.: Coccinellidae 

296, 299, 300Ð306, 312, 315, 316 

146, 150, 152, 153, 290, 292, 

Cryptolaemus wallacii Crotch Col.: Coccinellidae 153 

Cryptoprymna Hym.: Pteromalidae 243, 247 

Cryptus Hym.: Ichneumonidae 193 

Cryptus rutovinctus Pratt Hym.: Ichenumonidae 

culpator cincticornis, Stenichneumon 

curculionis, Physoderes 

curvicauda, Labia 

curvimaculatus, Chelonus 

curvispinosa, Crematogaster 

324 

Cuspicona simplex Walker Hem.: Pentatomidae 

custodiens, Anoplolepis 

225 

Cycloneda sanguinea Linnaeus Col.: Coccinellidae 332, 341 

Cydia ptychora, see Leguminivora ptychora 194 

Cylas formicarius (Fabricius) Col.: Curculionidae 

cylindrica, Sphaerophoria 

11 

Cylindromyia Dip.: Tachinidae 22


Cylindromyia rufifemur Paramonov Dip.: Tachinidae 202, 217, 233 

Cynipoide Hym.: Cynipidae 241, 247, 251 

Cyrtogaster Hym.: Pteromalidae 

cyrus, Telenomus 

245 

dactylopii, Leptomastix 

dactylopii, Prochiloneurus 

Dactylosternum abdominale (Fabricius) Col.: Hydrophilidae 85, 91, 93, 97, 102, 103 

Dactylosternum hydrophiloides Macleay Col.: Hydrophilidae 85, 91, 96Ð98, 100, 102 

Dactylosternum intermedium Reg. Col.: Hydrophilidae 91, 102 

Dactylosternum profundus Auct. Col.: Hydrophilidae 91, 102 

Dactylosternum subdepressum Lap. Col.: Hydrophilidae 91, 98 

Dactylosternum subquadratum Fairmaire Col.: Hydrophilidae 91 

darci, Alloxysta 

darvicola, Eurytoma 

dasyni, Anastatus 

dasyops, Winthemia 

Deanolis albizonalis (Hampson) Lep.: Pyralidae 110 

Deanolis sublimbalis Snellen Lep.: Pyralidae 2, 105Ð112 

deion, Trichogramma 

delhiensis, Lysiphlebus 

Delta pyriforme Fabricius Hym.: Eumenidae 25, 30 

deltiger, Toxares 

delucchii, Asecodes 

delucchii, Teleopterus 

demolitor, Microplitis 

Dendrocerus Hym.: Megaspilidae 65, 68 

dendrolimi, Trichogramma 

Dermatopelte Hym.: Eulophidae 191, 192 

Dermatopolle, see Dermatopelte 191 

derogata, Syllepte 

detorquens, Technomyrmex 

Diadegma Hym.: Ichneumonidae 325 

Diadegma insulare (Cresson) Hym.: Ichneumonidae 325, 348 

Diadegma plutellae Hym.: Ichneumonidae 325 

diadema, Sinea 

Diadiplosis hirticornis Felt Dip.: Cecidomyiidae 293 

Diaeretiella rapae (M'Intosh) Hym.: Braconidae 38, 48, 53


Diaphorencyrtus aligarhensis (Shafee, Alam and Agarwal) Hym.: Encyrtidae 113, 120, 

121, 123Ð133 

Diaphorencyrtus diaphorinae, see Diaphorencyrtus aligarhensis 121 

Diaphorina auberti Hollis Hem.: Psyllidae 116, 131 

Diaphorina cardiae Crawford Hem.: Psyllidae 131 

Diaphorina citri Kuwayama Hem.: Psyllidae 1, 113Ð134 

Diaretus rapae, see Diaeretiella rapae 

Diaspis bromeliae (Keiren) Hem.: Diaspididae 146, 153 

Dichomeris eridantis Meyrick Lep.: Gelechiidae 194 

Dichrodiplosis sp. Dip.: Cecidomyiidae 293 

Dicrodiplosis guatemalensis Felt Dip.: Cecidomyiidae 151 

dictynna, Aphanogmus 

dictynna, Calliceras 

dictynna, Ceraphron 

Dictyotis caenosus (Westwood) Hem.: Pentatomidae 225 

dilabida, Sturmia 

dimidiata, Harmonia 

dimidiata, Leis 

Dindymus rubiginosus (Fabricius) Hem.: Pyrrhocoridae 166, 168 

Diomus Col.: Coccinellidae 150, 300 

Diomus flavifrons see Diomus pumilio 

Diomus margipallens, see Scymnus margipallens 150 

Diomus pumilio (Weise) Col.: Coccinellidae 292, 306, 310, 313 

discolor, Micraspis 

Disophrys lutea (BrullŽ) Hym.: Braconidae 24, 30 

dispar, Lymantria 

Dissolcus fulviventris, see Gryon fulviventris 227 

diversus, Pheidologeton 

dolichostigma, Melanagromyza 

Dolicoderus Hym.: Formicidae 223 

dolosus, Enicospilus 

donacis, Melanaphis 

douglasi, Achrysocharis 

douglasi, Chrysonotomyia 

Drosica abjectella Walker Lep.: Tineidae 147, 154 

dryi, Tamarixia 

duplifascialis, Hendecasis 

Dysdercus Hem.: Pyrrhocoridae 135, 138


Dysdercus cingulatus (Fabricius) Hem.: Pyrrhocoridae 1, 2, 135Ð139 

dysmicocci, Pseudaphycus 

Dysmicoccus brevipes (Cockerell) Hem.: Pseudococcidae 1, 2, 141Ð156, 313 

Dysmicoccus neobrevipes Beardsley Hem.: Pseudococcidae 141Ð144, 152Ð154 

Earias Lep.: Noctuidae 28 

Earias biplaga Walker Lep.: Noctuidae 30 

Earias insulana Boisduval Lep.: Noctuidae 

eastopi, Trioza 

194 

Echthromorpha agrestoria (Swederus) Hym.: Ichneumonidae 25, 27 

Echthromorpha punctum Brutte Hym.: Ichneumonidae 325 

Ectophasia crassipennis (Fabricius) Dip.: Tachinidae 202, 224 

Ectophasiopsis arcuata (Bigot) Dip.: Tachinidae 202, 213, 221, 233, 324 

edentata, Voria 

Elachertus Hym.: Eulophidae 268, 273, 283 

Elasmus Hym.: Elasmidae 265, 273, 275 

Elasmus corbetti Ferri re Hym.: Elasmidae 265 

Elasmus tischeriae (Howard) Hym.: Elasmidae 265 

Elasmus zehntneri Ferri re Hym.: Elasmidae 

elegantilis, Neoleucinodes 

Elpe, see Camplyocheta 

265, 276, 284 

Encarsia Hym.: Aphelinidae 124, 125 

Encarsia shafeei, see Encarsia transvena 125 

Encarsia transvena (Timberlake) Hym.: Aphelinidae 125 

Endaphis maculans (Barnes) Dip.: Cecidomyiidae 57, 79 

Enicospilus Hym.: Ichneumonidae 25, 30, 325 

Enicospilus dolosus (Tosquinet) Hym.: Ichneumonidae 25, 28 

Enicospilus samoana (Kohl) Hym.: Ichneumonidae 25, 27 

Ephedrus Hym.: Aphidiidae 38, 71, 83 

Ephedrus cerasicola Starù Hym.: Aphidiidae 38, 67 

Ephedrus nacheri Quilis Hym.: Aphidiidae 38, 53, 64 

Ephedrus persicae Froggatt Hym.: Aphidiidae 38, 53, 64, 74, 83 

Ephedrus plagiator (Nees) Hym.: Aphidiidae 38, 53, 64, 71, 74, 75 

Epiclerus nomocerus Hym.: Tetracampidae 244 

Episyrphus balteatus (De Geer) Dip.: Syrphidae 

epius, Spalgis 

equatus, Trioxys 

43 

Eriborus argenteopilosus (Cameron) Hym.: Ichneumonidae 191, 194


eridantis, Dichomeris 

Eriococcus Hem.: Eriococcidae 312 

Eriosoma lanigerum (Hausmann) Hem.: Aphididae 

erosa, Anomis 

erosa, Cosmophila 

ervi, Aphidius 

erythraea, Chrysonotomyia 

erytreae, Trioza 

72 

Eublemma amabilis Moore Lep.: Noctuidae 194 

Euborellia annulipes (Lucas) Derm.: Labiidae 90, 102 

Euborellia pallipes (Shiraki) Derm.: Labiidae 22, 29 

Eucanthecona furcellata (Wolff) Hem.: Pentatomidae 22, 30 

Eucelatoria armigera (Coquillett) Dip.: Tachinidae 327, 328 

Eucelatoria rubentis (Coquillett) Dip.: Tachinidae 328 

Euclytia flava (Townsend) Dip.: Tachinidae 202 

Eucoilidea Hym.: Cynipidae 241, 247, 255 

Euderus Hym.: Eulophidae 241, 251, 268 

Eulissus Col.: Staphylinidae 92 

Eumenes campaniformis (Fabricius) Hym.: Eumenidae 25, 30 

Eumenes pyriformis, see Delta pyriforme 25, 30 

Eupelmus Hym.: Eupelmidae 242 

Eupelmus australiensis (Girault) Hym.: Eupelmidae 239, 242 

Eupelmus grayi Girault Hym.: Eupelmidae 242 

Eupelmus popa, see Eupelmus australiensis 239 

Eupelmus urozonus Dalman Hym.: Eupelmidae 242, 271 

Eupeodes confrater (Wiedemann) Dip.: Syrphidae 43 

Euphorocera Dip.: Tachinidae 328 

Euphorocera tachinomoides Townsend Dip.: Tachinidae 328 

Euplectrus Hym.: Eulophidae 324 

Euplectrus comstockii Howard Hym.: Eulophidae 324 

Euplectrus manilae Ashmead Hym.: Eulophidae 24 

Euplectrus platyhypenae Howard Hym.: Eulophidae 324, 337 

Eurydinotellus viridicoxa, see Callitula viridicoxa 251 

Euryrhopalus propinquus Kerrich Hym.: Encyrtidae 145, 147, 149 

Euryrhopalus schwarzi (Howard) Hym.: Encyrtidae 145, 147, 149 

Euryrophalus pretiosa, see Euryrophalus schwarzi 145, 147 

Eurytenes nanus, see Opius phaseoli 252 

Eurytoma Hym.: Eurytomidae 239, 242, 246, 247, 271, 277


Eurytoma larvicola Girault Hym.: Eurytomidae 242, 245 

Eurytoma poloni Girault Hym.: Eurytomidae 242, 247, 251 

Eurytoma syleptae Ferri re Hym.: Eurytomidae 28 

Eusandalum incompleta (Bou‹ek) Hym.: Eupelmidae 

euschisti, Trissolcus 

271, 274, 277 

Eutectona macheralis (Walker) Lep.: Pyralidae 194 

Euthyrhynchus floridanus (Linnaeus) Col.: Pentatomidae 330 

Eutochia pulla (Erichson) Col.: Tenebrionidae 92 

Eutrichopodopsis nitens, see Trichopoda giacomellii 202, 221, 232 

Euzophera ferticella Ragonot Lep.: Pyralidae 

evanescens, Trichogramma 

194 

Evania appendigaster (Linnaeus) Hym.: Evaniidae 

exaltatorius, Trogus 

exigua, Spodoptera 

exiguae, Hyposoter 

exiguum, Trichogramma 

111 

Exochomus flavipes (Thunberg) Col.: Coccinellidae 43, 292, 305, 310 

Exochomus flaviventris Mader Col.: Coccinellidae 292 

Exochomus metallicus Korsch Col.: Coccinellidae 

exoletum, Praon 

299, 310 

Exorista apicalia Baranov Dip.: Tachinidae 23 

Exorista sorbillans (Wiedemann) Dip.: Tachinidae 23, 27 

Exoristobia philippensis Ashmead Hym.: Encyrtidae 

exsaguis, Zelus 

201, 221 

fabae, Aphis 

fabarum, Lysiphlebus 

facialis, Pediobius 

fecundus, Ooencyrtus 

Ferrisia virgata Cockerell Hem.: Pseudococcidae 314 

ferrugatus, Belonuchus 

ferrugineus , Bactrocera 

ferruginosus, Chrysopilus 

ferticella, Euzophera 

ficus, Planococcus 

filicornis, Callitula 

flava, Anomis 

flava, Cosmophila


flava, Euclytia 

flavicincta, Campoletis 

flavipes, Aphelinus 

flavipes, Aphidius 

flavipes, Exochomus 

flavitarsis, Mischocyttarus 

flaviventris, Exochomus 

flavoorbitalis, Cremastus 

flavoorbitalis, Trathala 

flavopicta, Spilochalcis 

floridaensis, Aphidius 

floridanum, Copidosoma 

floridanus, Euthyrhynchus 

Fopius Hym.: Braconidae 240, 248, 251 

formicarius, Cylas 

formosa, Chrysonotomyia 

formosa, Chrysopa 

formosana, Schizobremia 

formosanus, Chelonus 

fragilis, Meteorus 

fragilis, Pseudococcus 

frauenfeldi, Bactrocera 

friedae, Camponotus 

fullawayi, Zaplatycerus 

Fulvius nigricornis Poppius Hem.: Miridae 90, 102 

fulviventris, Dissolcus 

fulviventris, Gryon 

fulviventris, Hadronotus 

funebris, Aphidius 

funeralis, Ascotolinx 

furcellata, Cantheconidia 

furcellata, Eucanthecona 

furnacalis, Ostrinia 

fuscifemora, Nepiera 

fuscipennis, Amblyteles 

galactopus, Baryscapus 

Galepsomyia Hym.: Eulophidae 268, 273, 275


Galerita bicolor (Drury) Col.: Carabidae 90, 102 

Gambrus ultimus (Cresson) Hym.: Ichneumonidae 325 

Gelis tenellus (Say) Hym.: Ichneumonidae 325 

geminata, Solenopsis 

Geocoris Hem.: Lygaeidae 22, 69 

Geocoris pallens StŒl Hem.: Lygaeidae 330, 340 

Geocoris punctipes Say Hem.: Lygaeidae 330 

Geotomus pygmaeus Dallas Hem.: Cydnidae 90, 102, 103 

gestuosus, Pterocormus 

giacomellii, Trichopoda 

gifuensis, Aphidius 

gifuensis, Telenomus 

gilva, Timberlakia 

Gitonides perspicax Knab Dip.: Drosophilidae 147, 153 

Glaucias amyoti (Dallas) Hem.: Pentatomidae 225 

globula, Hyperaspis 

glomerata, Cotesia 

Glyptapanteles vitripennis Curtis Hym.: Braconidae 336 

Goniozus Hym.: Bethylidae 166, 168, 182 

gossypii, Aphelinus 

gossypii, Aphis 

gracilis, Lipolexis 

graelsii, Xanthodes 

gramineum, Acrosternum 

graminis, Antonina 

graminum, Schizaphis 

grandis, Alcaeorrhynchus 

granulatus, Rogas 

grayi, Eupelmus 

greeni, Anagyrus 

greeni, Bracon 

gregori, Sympiesis 

Gryon Hym.: Scelionidae 201, 205, 228, 233 

Gryon antestiae, see Gryon fulviventris 227 

Gryon fulviventris (Crawford) Hym.: Scelionidae 205, 227 

Gryon japonicum (Ashmead) Hym.: Scelionidae 205, 212, 221 

Gryon obesum Masner Hym.: Scelionidae 205, 212, 221 

guatemalensis, Dicrodiplosis


guineense, Tetramorium 

gustavoi, Trichopoda 

Gymnosoma clavata (Rohdendorf) Dip.: Tachinidae 202, 233 

Gymnosoma kuramanum Matsumura Dip.: Tachinidae 202 

Gymnosoma rotundata (Fabricius) Dip.: Tachinidae 202, 233, 234 

Hadrojoppa cognatoria Smith Hym.: Ichneumonidae 12 

Hadronotus antestiae, see Gryon fulviventris 227 

Hadronotus fulviventris, see Gryon fulviventris 277 

Halticoptera Hym.: Pteromalidae 243, 247 

Halticoptera circulus (Walker) Hym.: Pteromalidae 243, 252 

Halticoptera patellana (Dalman) Hym.: Pteromalidae 243, 246, 249 

Hambletonia pseudococcina Compere Hym.: Encyrtidae 

hampei, Hypothenemus 

hampei, Stephanoderes 

hamygurivara, Sphegigaster 

hapaliae, Cremastus 

145, 148, 149, 152, 154Ð156 

Harmonia axyridis (Pallas) Col.: Coccinellidae 62, 63, 66, 128 

Harmonia dimidiata (Fabricius) Col.: Coccinellidae 43, 68 

Harmonia octomaculata (Fabricius) Col.: Coccinellidae 60, 292, 300 

Hebertia Hym.: Pteromalidae 243 

Helicoverpa Lep.: Noctuidae 27 

Helicoverpa armigera (HŸbner) Lep.: Noctuidae 

helva, Bogosia 

helymus, Metachaeta 

helymus, Periscepsia 

28, 30, 194 

Hemerobius Neu.: Hemerobiidae 331 

Hemiptarsenus Hym.: Eulophidae 241 

Hemiptarsenus semialbicornis, see Hemiptarsenus varicornis 252 

Hemiptarsenus varicornis (Girault) Hym.: Eulophidae 

hemipterus, Xenoencyrtus 

241, 252 

Hendecasis duplifascialis Hampson Lep.: Pyralidae 194 

Herbertia Hym.: Pteromalidae 239, 243 

Herse convolvuli, see Agrius convolvuli 10 

Hesperus sparsior (Bernhauer) Col.: Staphylinidae 92 

Heteropsylla cubana Crawford Hem.: Psyllidae 134 

Heteropsylla spinulosa Muddiman, Hodkinson & Hollis Hem.: Psyllidae 134


Heterospilus coffeicola Schmiedeknecht Hym.: Braconidae 

177, 178, 181, 182 

166, 169Ð171, 173, 175, 

Hexacladia hilaris Burks Hym.: Encyrtidae 

hilare, Acrosternum 

hilaris, Hexacladia 

hindecasisella, Phanerotoma 

203 

Hippodamia convergens (GuŽrin-MŽneville) Col.: Coccinellidae 69, 332, 341 

Hippodamia parenthesis (Say) Col.: Coccinellidae 332 

Hippodamia quinqesignata punctulata Le Conte Col.: Coccinellidae 332 

Hippodamia variegata (Goeze) Col.: Coccinellidae 

hirticornis, Diadiplosis 

62, 69 

Hister niloticus Marseul Col.: Histeridae 

hockiana, Carcelia 

hoffmanni, Scymnus 

hokkaidensis, Trioxys 

91 

Holcopelte Hym.: Eulophidae 268 

Hololepta Col.: Histeridae 91, 97, 98, 102 

Hololepta minuta Erichson Col.: Histeridae 97 

Hololepta quadridenta (Fabricius) Col.: Histeridae 91, 96Ð 98, 100, 101 

Hololepta striaditera Marseul Col.: Histeridae 91 

Homalotylus Hym.: Encyrtidae 306 

Horismenus Hym.: Eulophidae 268, 273 

Horismenus sardus Hym.: Eulophidae 

horticola, Chromatomyia 

howardi, Tetrastichus 

hullensis, Trissolcus 

humilis, Aphelinus 

humilis, Iridomyrmex 

268 

Hyalopterus pruni (Geoffroy) Hem.: Aphididae 

hybneri, Piezodorus 

hydrophiloides, Dactylosternum 

73 

Hypantropha Dip.: Tachinidae 328 

Hyperaspis Col.: Coccinellidae 150, 292, 300, 310 

Hyperaspis albicollis Gorham Col.: Coccinellidae 150 

Hyperaspis c-nigrum Mulsant Col.: Coccinellidae 150, 300 

Hyperaspis globula Casey Col.: Coccinellidae 310 

Hyperaspis jucunda Mulsant Col.: Coccinellidae 310 

Hyperaspis lateralis Mulsant Col.: Coccinellidae 292


Hyperaspis polita Weise Col.: Coccinellidae 292, 306 

Hyperaspis silvestrii Weise Col.: Coccinellidae 150, 154 

Hyperomyzus lactucae (Linnaeus) Hem.: Aphididae 62 

Hyphantrophaga Dip.: Tachinidae 327 

Hyposolenus laevigatus, see Plaesius laevigatus 91 

Hyposoter exiguae (Viereck) Hym.: Ichneumonidae 317, 325, 338, 340Ð343, 346, 347 

Hypothenemus Col.: Scolytidae 163, 164, 178 

Hypothenemus hampei (Ferrari) Col.: Scolytidae 1, 2, 157Ð183 

illota, Carcelia 

importatus, Opius 

includens, Chrysodeixis 

includens, Nephus 

includens, Pseudoplusia 

includens, Scymnus 

incompleta, Eusandalum 

incompleta, Ratzeburgiola 

inconspicua, Palexorista 

indagator, Sycanus 

indefensa, Plutarchia 

indica, Cosmophila 

indicum, Monomorium 

indicus, Paragus 

indicus, Trioxys 

infesta, Casinaria 

ingenuus, Cirrospilus 

innota, Sarcodexia 

insidiosus, Orius 

insulana, Earias 

insulare, Diadegma 

insularis, Angitia 

insularis, Blepyrus 

insularis, Chelonus 

insularis, Omicrogiton 

intacta, Thopeutis 

intermedia, Brachymeria 

intermedium, Dactylosternum 

interocularis, Thyreocephalus


interpunctella, Plodia 

intima, Chrysopa 

Iridomyrmex humilis Mayr Hym.: Formicidae 

irregularis, Chrysopa 

144 

Ischiodon aegyptius (Wiedemann) Dip.: Syrphidae 43 

Ischiodon scutellaris Fabricius Dip.: Syrphidae 43, 68 

Iseropus stercorator orgyiae (Ashmead) Hym.: Ichneumonidae 325 

Isyropa Dip.: Tachinidae 22 

Itamoplex Hym.: Ichneumonidae 191Ð193 

Itoplectis conquisator (Say) Hym.: Ichneumonidae 325 

japonica, Lysiphlebia 

japonica, Propylea 

japonicum, Gryon 

japonicum, Trichogramma 

japonicus, Anastatus 

japonicus, Meteorus 

japonicus, Pleurotropposis 

japonicus, Stenomesius 

javanus, Plaesius 

johnsoni, Ooencyrtus 

jokahamae, Polistes 

josefi, Clausenia 

jucunda, Hyperaspis 

kamburovi, Allotropa 

kashmiriensis, Aphelinus 

kenyae, Planococcus 

kivuensis, Anagyrus 

koebelei, Cleodiplosis 

Kratoysma Hym.: Eulophidae 269, 272, 277 

Kratoysma citri Bou‹ek Hym.: Eulophidae 269, 276, 284 

kraunhiae, Planococcus 

kulingensis, Chrysopa 

kuramanum, Gymnosoma 

Labia borellii Burr Derm.: Labiidae 90 

Labia curvicauda (Motschulsky) Derm.: Labiidae 90


Labidura riparia Pallas Derm.: Labiduridae 330 

lacciperda, Chrysopa 

lacciperda, Odontochrysa 

lacciperda, Plesiochrysa 

lactucae, Hyperomyzus 

lacunatus, Psix 

laeviceps, Cotesia 

laevigatus, Hyposolenus 

laevigatus, Plaesius 

lanata, Chrysopa 

lanigerum, Eriosoma 

lanipes, Trichopoda 

laphygmae, Meteorus 

larvicola, Eurytoma 

lasus, Brachymeria 

lateralis, Hyperaspis 

Lebia analis Dejean Col.: Carabidae 332 

Leguminivora ptychora (Meyrick) Lep.: Tortricidae 194 

leibyi, Schizocerophaga 

Leis dimidiata, see Harmonia dimidiata 

leopardina, Marietta 

lepelleyi, Trissolcus 

Leptochirus unicolor, see Priochirus unicolor 92, 93, 102 

Leptomastidea abnormis (Girault) Hym.: Encyrtidae 287, 295Ð297, 299Ð301, 303Ð307, 

313, 315, 316 

Leptomastix Hym.: Encyrtidae 301 

Leptomastix dactylopii Howard Hym.: Encyrtidae 148, 149, 287, 295Ð297, 299Ð307, 

312Ð316 

Leptomastix nigrocoxalis Compere Hym.: Encyrtidae 295, 297 

Leptomastix trilongifasciatus Girault Hym.: Encyrtidae 295, 297 

Lespesia sp. Dip.: Tachinidae 328 

Lespesia achaetoneura Dip.: Tachinidae 328 

Lespesia archippivora (Riley) Dip.: Tachinidae 328 

Leucinodes orbonalis GuenŽe Lep.: Pyralidae 1, 2, 185Ð195 

Leucopis Dip.: Chamaemyiidae 43, 65 

Leucopis alticeps Czerny Dip.: Chamaemyiidae 293, 306 

Leucopis bella Loew Dip.: Chamaemyiidae 293 

Leucopis silesiaca Eggar Dip.: Chamaemyiidae 293


lilacinus, Planococcus 

lineatus, Brumoides 

Lioderma Col.: Histeridae 91, 102 

Lioderma quadridentata, see Hololepta quadridentata 91 

liogaster, Opius 

Lipolexis ambiguus, see Lysiphlebus ambiguus 39 

Lipolexis gracilis Fšrster Hym.: Aphidiidae 38, 53, 68, 75 

Lipolexis pseudoscutellaris, see Lipolexis scutellaris 54 

Lipolexis scutellaris Mackauer Hym.: Aphidiidae 39, 54, 64, 66Ð68, 75, 82, 83 

Liriomyza brassicae (Riley) Dip.: Agromyzidae 252 

Liriomyza sativae Blanchard Dip.: Agromyzidae 252 

Liriomyza trifolii (Burgess) Dip.: Agromyzidae 251 

liriomyzae, Meruana 

Lissauchenius venator LafertŽ Col.: Carabidae 26 

littoralis, Spodoptera 

lituratis, Piezodorus 

Lixophaga Dip.: Tachinidae 195 

Lobodiplosis pseudococci, see Vincentodiplosis pseudococci 151 

lodosi, Trissolcus 

longefasciatus, Cirrospilus 

longicornis, Paratrechina 

longipes, Anoplolepis 

longispinus, Pseudococcus 

longiventris, Paragus 

lonicerae, Aphidius 

louisianae, Scymnus 

lucens, Asaphes 

lunata, Cheilomenes 

lurida, Scotinophara 

lutea, Disophrys 

lutea, Oligochrysa 

lutea, Ophelosia 

Lygocerus, see Dendrocerus 65 

Lymantria dispar (Linnaeus) Lep.: Lymantriidae 225 

lyncus, Cirrospilus 39 

Lysaphidus Hym.: Aphidiidae 37 

Lysaphidus platensis, see Aphidius colemani 37 

Lysaphidus schimitscheki (Fahriner) Hym.: Aphidiidae 54


Lysiphlebia Hym.: Aphidiidae 39 

Lysiphlebia japonica (Ashmead) Hym.: Aphidiidae 39, 54, 63, 64, 66, 75, 82, 83 

Lysiphlebia mirzai Shuja Uddin Hym.: Aphidiidae 54, 70, 75 

Lysiphlebia rugosa Starù & Schlinger Hym.: Aphidiidae 39 

Lysiphlebus Hym.: Aphidiidae 40, 55, 63 

Lysiphlebus ambiguus, see Lysiphlebus fabarum 39, 54, 71 

Lysiphlebus cardui, see Lysiphlebus fabarum 54 

Lysiphlebus confusus, see Lysiphlebus fabarum 39, 54, 75 

Lysiphlebus delhiensis (Subba Rao & Sharma) Hym.: Aphidiidae 39 

Lysiphlebus fabarum (Marshall) Hym.: Aphidiidae 39, 54, 60, 61, 65Ð67, 71, 75, 83 

Lysiphlebus salicaphis, see Adialytus salicaphis 37 

Lysiphlebus shaanxiensis Chou and Xian Hym.: Aphidiidae 54 

Lysiphlebus testaceipes (Cresson) Hym.: Aphidiidae 

76, 77, 82, 83 

5, 40, 55, 60, 61, 63, 64, 66Ð69, 74, 

macheralis, Eutectona 

macheralis, Pyrausta 

macrophallus, Sturmia 

macrosiphophagum, Toxares 

maculans, Endaphis 

maculata fuscilabris, Ceratomegilla 

maculipennis, Pseudaphycus 

maculiventris, Podisus 

madecassa, Brachymeria 

Madremyia saundersii (Williston) Dip.: Tachinidae 328 

maidis, Rhopalosiphum 

malayensis, Ooencyrtus 

mali, Aphelinus 

Mallada basalis (Walker) Neu.: Chrysopidae 291, 302 

Mallada boninensis Okamoto Neu.: Chrysopidae 271, 291, 302 

manilae, Euplectrus 

manilae, Snellenius 

marginata, Mesograpta 

marginatum, Acrosternum 

marginellus, Collops 

marginipallens, Diomus 

marginiventris, Cotesia 

margipallens, Scymnus


mariae, Spilochalcis 

Marietta Hym.: Aphelinidae 128 

Marietta javensis, see Marietta leopardina 124, 129 

Marietta leopardina Motschulsky Hym.: Aphelinidae 124, 126, 149 

mariscusae, Aphelinus 

maro, Trissolcus 

marshalli, Sericophoromyia 

Maruca testulalis, see Maruca vitrata 194 

Maruca vitrata Fabricius Lep.: Pyralidae 194 

matricariae, Aphidius 

mauritiusi, Scymnus 

mecrida, Allotropa 

mediterraneus, Carpocoris 

mediterraneus, Pnigalio 

megacephala, Pheidole 

megacephala, Pheidole 

Megaselia rufipes (Meigen) Dip.: Phoridae 12, 15 

Melanagromyza Dip.: Agromyzidae 236 

Melanagromyza atomella (Malloch) Dip.: Agromyzidae 253 

Melanagromyza dolichostigma de Meijere Dip.: Agromyzidae 236, 247, 251, 254 

Melanagromyza obtusa (Malloch) Dip.: Agromyzidae 236 

Melanagromyza phaseoli, see Ophiomyia phaseoli 

Melanagromyza sojae (Zehnter) Dip.: Agromyzidae 236, 247, 248, 251, 253, 254 

melanagromyzae, Biosteres 

melanagromyzae, Opius 

Melanaphis donacis (Passerini) Hem.: Aphididae 73 

melvillei, Tropidophryne 

Menochilus sexmaculatus, see Cheilomenes sexmaculata 65 

menthastri, Sphaerophoria 

Merismorella shakespearei, see Syntomopus shakespearei 254 

Meruana liriomyzae Bou‹ek Hym.: Eulophidae 241, 252 

Mesochorus Hym.: Ichneumonidae 25, 29 

Mesochorus africanus (Ferri re) Hym.: Ichneumonidae 28 

Mesograpta marginata (Say) Dip.: Syrphidae 332 

Metachaeta helymus Walker Dip.: Tachinidae 328 

metallicus, Exochomus 

Meteorus Hym.: Braconidae 21 

Meteorus autographae Muesebeck. Hym.: Braconidae 322, 348


Meteorus fragilis Wesmael Hym.: Braconidae 24, 29 

Meteorus japonicus, see Meteorus pulchricornis 24 

Meteorus laphygmae Viereck Hym.: Braconidae 323 

Meteorus pulchricornis (Wesmael) Hym.: Braconidae 21, 24, 29 

Metopius Hym.: Ichneumonidae 

metricus, Polistes 

25 

Micraspis discolor (Fabricius) Col.: Coccinellidae 65 

Microbracon phyllocnistidis, see Bracon phyllocnistidis 265, 274 

Microcharops bimaculata (Ashmead) Hym.: Ichneumonidae 325, 335 

Microcharops tibialis (Cresson) Hym.: Ichneumonidae 325 

Microdon bicolor Sack Dip.: Syrphidae 43 

Microgaster brassicae (Muesebeck) Hym.: Braconidae 

347 

317, 323, 338Ð341, 343, 346, 

Microgaster demolitor, see Microplitis demolitor 336 

Microgaster plutellae (Muesebeck) Hym.: Braconidae 323 

Microgaster rufiventris, see Microplitis rufiventris 356 

Micromus posticus (Walker) Neu.: Hemerobiidae 69 

Micromus timidus (Fabricius) Neu.: Hemerobiidae 68 

Microplitis alaskensis Ashmead Hym.: Braconidae 323 

Microplitis brassicae, see Microgaster brassicae 323 

Microplitis demolitor Wilkinson Hym.: Braconidae 356 

Microplitis rufiventris Kokujev Hym.: Braconidae 

mikan, Sympiesomorpha 

minio, Pnigalio 

minuta, Hololepta 

minutum, Trichogramma 

minutus, Orius 

mirzai, Lysiphlebia 

356 

Mischocyttarus flavitarsis (Saussure) Hym.: Vespidae 

mitsukurii, Asolcus 

mitsukurii, Trissolcus 

molestus, Rogas 

333 

Monomorium Hym.: Formicidae 153 

Monomorium indicum Forel Hym.: Myrmecidae 

montana, Winthemia 

montanus, Patrocloides 

montrouzieri, Cryptolaemus 

mormideae, Telenomus 

67


multicolor, Brachymeria 

multilineatum, Zagrammosoma 

murrayii, Psylla 

myzophagum, Praon 

Myzus Hem.: Aphididae 75 

Myzus persicae Sulzer Hem.: Aphididae 5, 59, 63, 71Ð74 

Nabis alternatus Parshley Col.: Nabidae 330 

Nabis americoferus Carayon Hem.: Nabidae 330, 340 

Nabis roseipennis (Reuter) Hem.: Nabidae 330 

Nabis sinoferus Hsiao Hem.: Nabidae 22 

nacheri, Ephedrus 

nakagawai, Telenomus 

nanus, Eurytenes 

narangae, Zacharops 

nasalis, Cermatulus 

nasuta, Allograpta 

nasuta, Prorops 

nearctaphidis, Trioxys 

nebulosa, Pseudiastata 

neobrevipes, Dysmicoccus 

Neochrysocharis, see Chrysonotomyia 266, 277 

Neodimmockia agromyzae, see Hemiptarsenus varicornis 252 

Neoleucinodes Lep.: Pyralidae 186 

Neoleucinodes elegantilis (GuenŽe) Lep.: Pyralidae 186, 195 

Neoprochiloneurus bolivari (Mercet) Hym.: Encyrtidae 306 

Neorileya Hym.: Eurytomidae 203 

nepalensis, Cristicaudus 

Nephopterix rhodobasalis Hampson Lep.: Pyralidae 194 

Nephus Col.: Coccinellidae 300, 310 

Nephus bilucenarius, see Scymnus bilucenarius 145, 150 

Nephus bipunctatus (Kugelann) Col.: Coccinellidae 292, 298, 306, 310 

Nephus includens, see Scymnus includens 292 

Nephus pictus, see Scymnus bilucenarius 145 

Nephus reunioni (Fursch) Col.: Coccinellidae 292, 302, 304, 306, 310, 316 

Nepiera fuscifemora Graf Hym.: Ichneumonidae 326 

nerii, Aphis 

Nesolynx phaeosoma (Waterston) Hym.: Eulophidae 28 

Netelia Hym.: Ichneumonidae 325


Nezara Hem.: Pentatomidae 198, 202 

Nezara antennata Scott Hem.: Pentatomidae 208, 224, 229, 233, 234 

Nezara viridula (Linnaeus) Hem.: Pentatomidae 1, 137, 197Ð233 

nezarae, Ooencyrtus 

ni, Trichoplusia 

niger, Asaphoideus 

niger, Thysanus 

niger, Xenoencyrtus 

nigriceps, Cardiochiles 

nigricornis, Chrysopa 

nigricornis, Fulvius 

nigrifrontalis, Trichopoda 

nigripes, Sigalphus 

nigritus, Aphelinus 

nigrocoxalis, Leptomastix 

nigrofasciatus, Phonoctonus 

nigronervosa, Pentalonia 

niloticus, Hister 

nitens, Eutrichopodopsis 

noctuae, Zenillia 

nomocerus, Epiclerus 

Noorda albizonalis, see Deanolis albizonalis 106 

Norbanus Hym.: Pteromalidae 239 

notata, Sympiesis 

novemnotata francisciana, Coccinella 

Nyereria Hym.: Braconidae 24, 28 

obesum, Gryon 

obscurata, Brachymeria 

obscurus, Pseudococcus 

obtusa, Agromyza 

obtusa, Melanagromyza 

octomaculata, Harmonia 

octopunctata, Ankylopteryx 

Ocyptera, see Cylindromyia 42 

Odontochrysa lacciperda Kimmins Neu.: Chrysopidae 291, 302, 314 

Oechalia schellembergii (GuŽrin-MŽneville) Hem.: Pentatomidae 22, 27, 218 

oenone, Trissolcus


ogyges, Trissolcus 

oleracei, Opius 

Oligochrysa lutea (Walker) Neu.: Chrysopidae 291, 300 

Olla v-nigrum (Mulsant) Col.: Coccinellidae 332 

Omicrogiton insularis Orchym. Col.: Hydrophilidae 91, 102 

Ooencyrtus Hym.: Encyrtidae 201, 204, 223, 225, 230 

Ooencyrtus californicus Hym.: Encyrtidae 203 

Ooencyrtus fecundus Ferri re & VoegelŽ Hym.: Encyrtidae 204 

Ooencyrtus johnsoni (Howard) Hym.: Encyrtidae 204 

Ooencyrtus malayensis Ferri re Hym.: Encyrtidae 204 

Ooencyrtus nezarae Ishii Hym.: Encyrtidae 204, 212, 221, 224, 227 

Ooencyrtus pityocampae (Mercet) Hym.: Encyrtidae 204 

Ooencyrtus submetallicus (Howard) Hym.: Encyrtidae 204, 212Ð214, 217, 221, 227, 230 

Ooencyrtus trinidadensis Crawford Hym.: Encyrtidae 

Ophelosia crawfordi Riley Hym.: Encyrtidae 295, 298 

204, 213, 221 

Ophiomyia centrosematis (de Meijere) Dip.: Agromyzidae 239, 248, 252 

Ophiomyia phaseoli (Tryon) Dip.: Agromyzidae 1, 2, 235Ð255 

Ophiomyia spencerella Greathead Dip.: Agromyzidae 236, 239, 253, 255 

Opius Hym.: Braconidae 240, 247 

Opius importatus Fischer Hym.: Braconidae 235, 239, 240, 249, 250, 255 

Opius liogaster SzŽpligeti Hym.: Braconidae 240, 248, 252 

Opius melanagromyzae, see Opius phaseoli 252 

Opius oleracei Fischer Hym.: Braconidae 239, 240 

Opius phaseoli Fischer Hym.: Braconidae 

optimus, Abacetus 

orbana, Antestis 

orbonalis, Leucinodes 

orestes, Chrysopa 

235, 239, 240, 246, 248Ð250, 252, 253, 255 

Orius Hem.: Anthocoridae 43 

Orius insidiosus Say Col.: Anthocoridae 330 

Orius minutus (Linnaeus) Hem.: Anthocoridae 22, 63, 271, 291 

Orius tristicolor (White) Hem.: Anthocoridae 330, 340, 341 

Ostrinia furnacalis (GuenŽe) Lep.: Pyralidae 

ovata, Brachymeria 

14 

Oxyharma subaenea (Dodd) Hym.: Pteromalidae 243, 254 

Pachyneuron Hym.: Pteromalidae 67, 77 

Pachyneuron aphidis BouchŽ Hym.: Pteromalidae 57, 65, 68, 78


Pachyneuron concolor (Fšrster) Hym.: Pteromalidae 123, 125 

Pachyneuron solitarius (Ratz) Hym.: Pteromalidae 306 

Pachyneuron vitodurense Delucchi Hym.: Pteromalidae 68 

Pachyophthalmus Dip.: Tachinidae 

pacificus, Planococcus 

pacificus, Telenomus 

190 

Paederus Col.: Staphylinidae 223 

Palexorista inconspicua (Meigen) Dip.: Tachinidae 23 

Palexorista quadrizonula (Thomson) Dip.: Tachinidae 

pallens, Chrysopa 

pallens, Geocoris 

pallidicollis, Pullus 

pallidus, Trioxys 

pallipes, Euborellia 

paolii, Brachymeria 

parachrysops, Sturmia 

23, 30, 31 

Paragus bicolor, see Microdon bicolor 43 

Paragus borbonicus Macquart Dip.: Syrphidae 43 

Paragus indicus, see Paragus tibialis 43 

Paragus longiventris Loewe Dip.: Syrphidae 43 

Paragus serratus (Fabricius) Dip.: Syrphidae 43 

Paragus tibialis (FallŽn) Dip.: Syrphidae 

paramali, Aphelinus 

43 

Paranaemia vittegera (Mulsant) Col.: Coccinellidae 332 

Parapanteles Hym.: Braconidae 24, 28 

Paratrechina longicornis (Latreille) Hym.: Formicidae 145 

Paratrigonogastra stella, see Sphegigaster stella 

parenthesis, Hippodamia 

patellana, Halticoptera 

247, 254 

Patrocloides montanus (Cresson) Hym.: Ichneumonidae 325, 340 

Pauesia antennata Makerjl Hym.: Aphidiidae 40 

Pauridia peregrina, see Coccidoxenoides peregrinus 294, 297, 311 

Pediobius Hym.: Eulophidae 239, 241 

Pediobius acantha (Walker) Hym.: Eulophidae 241 

Pediobius facialis (Giraud) Hym.: Eulophidae 324, 336 

Pediobius sexdentatus Hym.: Pteromalidae 

pennipes, Trichopoda 

pennsylvanicum, Acrosternum 

326, 338


pensylvanica, Vespula 

Pentalonia nigronervosa Coquerel Hem.: Aphididae 67, 68, 74, 81 

pentheus, Chrysocharis 

perdignus, Pseudaphycus 

peregrina, Pauridia 

peregrinator, Calosoma 

peregrinus, Coccidoxenoides 

Periscepsia helymus, see Metachaeta helymus 327 

perla, Chrysopa 

perplexus, Rogas 

persicae, Ephedrus 

persicae, Myzus 

perspicax, Gitonides 

perticella, Euzophera 

petiolatus, Semielacher 

pfeifferi, Allograpta 

phaeosoma, Nesolynx 

Phanerotoma Hym.: Braconidae 190, 192, 195 

Phanerotoma hindecasisella Cameron Hym.: Braconidae 190, 194 

phaseoli, Agromyza 

phaseoli, Melanagromyza 

phaseoli, Ophiomyia 

phaseoli, Opius 

Pheidole Hym.: Formicidae 67, 153, 156 

Pheidole megacephala (Fabricius) Hym.: Formicidae 92, 93, 144, 152Ð154, 223, 300 

Pheidologeton diversus (Jerdon) Hym.: Formicidae 155 

Phenacoccus solani Ferris Hem.: Pseudococcidae 313 

philippensis, Exoristobia 

Phonoctonus Hem.: Reduviidae 139 

Phonoctonus nigrofasciatus StŒl Hem.: Reduviidae 139 

Phonoctonus subimpictus StŒl Hem.: Reduviidae 139 

Phorocera Dip.: Tachinidae 328 

phorodontis, Aphidius 

Phorticus pygmaeus Poppius Hem.: Nabidae 90, 102 

Phryxe vulgaris (FallŽn) Dip.: Tachinidae 328 

phyllocnistidis, Bracon 

phyllocnistidis, Microbracon 

Phyllocnistis Lep.: Gracillariidae 258


Phyllocnistis citrella Stainton Lep.: Gracillariidae 1, 2, 257Ð286 

phyllocnistis, Cirrospilus 

phyllocnistis, Scotolinx 

phyllocnistoides, Cirrospiloideus 

phyllocnistoides, Cirrospilus 

phyllocnistoides, Cirrostichus 

phyllocnistoides, Tetrastichus 

Phymastichus coffea La Salle Hym.: Eulophidae 157, 166, 168, 174, 178, 182 

Physoderes curculionis China Hem.: Reduviidae 90, 102 

Phytomyza albiceps Meigen Dip.: Agromyzidae 254 

Phytomyza atricornis, see Chromatomyia horticola 239, 252 

picipes, Aphidius 

pictus, Cirrospilus 

pictus, Nephus 

pictus, Scymnus 

Pieris rapae Linnaeus Lep.: Pieridae 337 

Piezodorus hybneri (Gmelin) Hem.: Pentatomidae 218, 227 

Piezodorus lituratus (Fabricius) Hem.: Pentatomidae 224 

pilipes, Trichopoda 

Pimpla aequalis (Provancher) Hym.: Ichneumonidae 325 

pinguis, Winthemia 

ÔPireniniÕ Hym.: Pteromalidae 276 

pisum, Acyrthosiphon 

pityocampae, Ooencyrtus 

Plaesius javanus Erichson Col.: Histeridae 85, 91, 93, 95Ð100, 103, 104 

Plaesius laevigatus Marseul Col.: Histeridae 85, 91, 93, 95, 96, 100, 104 

plagiator, Ephedrus 

Planococcus Hem.: Pseudococcidae 288 

Planococcus citri (Risso) Hem.: Pseudococcidae 1, 2, 155, 287Ð316 

Planococcus ficus (Signoret) Hem.: Pseudococcidae 288 

Planococcus kenyae (Le Pelley) Hem.: Pseudococcidae 288, 311 

Planococcus kraunhiae (Kuwana) Hem.: Pseudococcidae 296, 305 

Planococcus lilacinus Cockerell Hem.: Pseudococcidae 313, 314 

Planococcus pacificus Cox Hem.: Pseudococcidae 288, 313 

platensis, Aphidius 

platensis, Lysaphidius 

platneri, Trichogramma 

platyhypenae, Euplectrus


Platynaspis Col.: Coccinellidae 310 

Platynaspis capicola Crotch Col.: Coccinellidae 68 

Platysoma Col.: Histeridae 91 

Platysoma abrupta Erichson Col.: Histeridae 91, 102 

Plautia brunnipennis (Montrouzier and Signoret) Hem.: Pentatomidae 225 

Plautia crossota (Dallas) Hem.: Pentatomidae 206 

Plesiochrysa lacciperda, see Chrysopa lacciperda 

pleuralis, Alloxysta 

291 

Pleurotropitiella albipes Blanchard Hym.: Eulophidae 203 

Pleurotropposis japonicus (Kamijo) Hym.: Eulophidae 269 

Plodia interpunctella (HŸbner) Lep.: Pyralidae 

plusiae, Siphona 

344 

Plutarchia Hym.: Eurytomidae 242, 247 

Plutarchia bicarinativentris Girault Hym.: Eurytomidae 239, 242 

Plutarchia indefensa (Walker) Hym.: Eurytomidae 242, 248, 254 

Plutella xylostella Linnaeus Lep.: Yponomeutidae 

plutellae, Angitia 

plutellae, Cotesia 

plutellae, Diadegma 

plutellae, Microgaster 

337 

Pnigalio Hym.: Eulophidae 269, 273, 274, 277, 278 

Pnigalio agraules Walker Hym.: Eulophidae 269, 276 

Pnigalio mediterraneus, see Pnigalio agraules 276 

Pnigalio minio (Walker) Hym.: Eulophidae 

podisi, Telenomus 

269, 277, 284 

Podisus Hem.: Pentatomidae 220 

Podisus maculiventris (Say) Hem.: Pentatomidae 27, 331 

Polistes Hym.: Vespidae 26, 29 

Polistes apachus Saussure Hym.: Vespidae 333 

Polistes jokahamae Radoszkowski Hym.: Vespidae 26, 29 

Polistes metricus Say Hym.: Vespidae 

polita, Hyperaspis 

poloni, Eurytoma 

333 

Polycystomyia beneficia, see Callitula viridicoxa 251 

Polycystus Hym.: Pteromalidae 243, 246 

Polycystus propinquus Waterston Hym.: Pteromalidae 

popa, Eupelmus 

posticus, Micromus 

243


Praon Hym.: Braconidae 40, 56, 77 

Praon abjectum (Haliday) Hym.: Braconidae 40, 55 

Praon absinthii Bagnall Hym.: Braconidae 55, 64 

Praon exsoletum (Nees) Hym.: Braconidae 40, 55 

Praon myzophagum Mackauer Hym.: Braconidae 56 

Praon volucre (Haliday) Hym.: Braconidae 

pretiosa, Euryrophalus 

pretiosum, Trichogramma 

41, 56, 62, 77 

Priochirus unicolor (Laporte) Col.: Staphylinidae 92, 93, 102 

Pristomerus Hym.: Ichneumonidae 325 

Pristomerus spinator (Fabricius) Hym.: Ichneumonidae 325 

Pristomerus testaceus Morley Hym.: Ichneumonidae 191, 194 

Prochiloneurus dactylopii (Howard) Hym.: Encyrtidae 

profundum, Dactylosternum 

305 

Propagalerita bicolor, see Galerita bicolor 

propinquus, Euryrhopalus 

propinquus, Polycystus 

90 

Propylea japonica (Thunberg) Col.: Coccinellidae 62, 63 

Prorops nasuta Waterston Hym.: Bethylidae 157, 166, 168, 170Ð182 

Prospalta capensis, see Condica capensis 194 

Protomicroplitis Hym.: Braconidae 

pruinosus, Cosmopolites 

pruni, Hyalopterus 

24, 28 

Psalis americana, see Carcinophora americana 90, 102 

Pseudaphycus Hym.: Encyrtidae 148, 149, 152, 300 

Pseudaphycus angelicus (Howard) Hym.: Encyrtidae 152, 295, 298 

Pseudaphycus angustifrons Gahan Hym.: Encyrtidae 148 

Pseudaphycus dysmicocci Bennett Hym.: Encyrtidae 148, 149, 155 

Pseudaphycus maculipennis Mercet Hym.: Encyrtidae 295, 298, 314 

Pseudaphycus perdignus Compere and Zinna Hym.: Encyrtidae 295, 300, 301, 309 

Pseudiastata nebulosa Coquerell Dip.: Drosophilidae 147, 151, 152, 155, 156 

Pseudiastata pseudococcivora Sabrosky Dip.: Drosophilidae 

pseudococci, Anagyrus 

pseudococci, Lobodiplosis 

pseudococci, Vincentodiplosis 

pseudococcina, Hambletonia 

pseudococcivora, Pseudiastata 

151 

Pseudococcus brevipes, see Dysmicoccus brevipes 142


Pseudococcus citriculus Green Hem.: Pseudococcidae 303, 311 

Pseudococcus comstocki Kuwana Hem.: Pseudococcidae 313 

Pseudococcus fragilis Brain Hem.: Pseudococcidae 311 

Pseudococcus longispinus Targioni-Tozzetti Hem.: Pseudococcidae 311 

Pseudococcus obscurus Essig Hem.: Pseudococcidae 311 

Pseudococcus vitis Niediel Hem.: Pseudococcidae 

pseudomagnoliarum, Coccus 

313 

Pseudoperichaeta Dip.: Tachinidae 190 

Pseudoplusia includens, see Chrysodeixis includens 

pseudoscutellaris, Lipolexis 

psidii, Chloropulvinaria 

340 

Psix lacunatus Johnson & Masner Hym.: Scelionidae 205 

Psix striaticeps (Dodd) Hym.: Scelionidae 205, 220 

Psylla Hem.: Psyllidae 131 

Psylla citricola Yang and Li Hem.: Psyllidae 116 

Psylla citrisuga Yang and Li Hem.: Psyllidae 116 

Psylla murrayii Mathur Hem.: Psyllidae 116 

Psyllaephagus Hym.: Encyrtidae 123, 125, 129 

Psyllaephagus diaphorinae, see Diaphorencyrtus aligarhensis 121 

Pterocormus gestuosus (Cresson) Hym.: Ichneumonidae 326 

Pteromalus Hym.: Pteromalidae 205 

Pterosema subaenea, see Oxyharma subaenea 

ptychora, Cydia 

ptychora, Leguminivora 

pulchricornis, Meteorus 

pulla, Eutochia 

254 

Pullus Col.: Coccinellidae 43 

Pullus pallidicollis Mulsant Col.: Coccinellidae 292, 302 

Pulvinaria Hem.: Coccidae 

pumilio, Diomus 

punctata, Xanthopimpla 

punctipes, Geocoris 

punctum, Echthromorpha 

punicae, Aphis 

purpurea, Clausenia 

pygmaeus, Geotomus 

pygmaeus, Phorticus 

312 

Pyrausta macheralis, see Eutectona macheralis 194


pyreformis, Eumenes 

pyriforme, Delta 

pyrrhopya, Winthemia 

Quadrastichus Hym.: Eulophidae 269, 272, 275, 277, 281 

quadratus, Belonuchus 

quadridenta, Hololepta 

quadridentata, Lioderma 

quadripustulata, Winthemia 

quadristriata, Scotolinx 

quadristriatus, Cirrospilus 

quadrivittatus, Scymnus 

quadrizonula, Palexorista 

quatrosignata, Sticholotis 

quinqesignata punctulata, Hippodamia 

radiata, Tamarixia 

radiatus, Tetrastichus 

ramburi, Chrysopa 

rapae, Diaeretiella 

rapae, Diaretus 

rapae, Pieris 

Ratzeburgiola incompleta, see Eusandalum incompleta 271, 277 

remota, Bessa 

renardii, Zelus 

repanda, Coccinella 

reunioni, Nephus 

reunioni, Scymnus 

rhanis, Synopeas 

Rhizobius ventralis Erichson Col.: Coccinellidae 146 

rhodobasalis, Nephopterix 

Rhopalosiphum maidis (Fitch) Hem.: Aphididae 77 

Rhychium attrisium Van der Vecht Hym.: Vespidae 111 

ribis, Aphidius 

rietscheli, Trioxys 

riparia, Labidura 

Riptortus serripes (Fabricius) Hem.: Alydidae 218 

robusta, Sphaerophoria

oepkei, Scymnus 

Rogas Hym.: Braconidae 

Rogas granulatus De Gant Hym.: Braconidae 323 

Rogas molestus Cresson Hym.: Braconidae 323 

Rogas rufocoxalis Gahan Hym.: Braconidae 323 

rosae, Aphidius 

roseipennis, Nabis 

rotundata, Gymnosoma 

rubentis, Eucelatoria 

rubicola, Trioxys 

rubiginosus, Dindymus 

rubricatus, Xenoencyrtus 

rudus, Trissolcus 

ruficrus, Apanteles 

ruficrus, Cotesia 

rufifemur, Cylindromyia 

rufilabris, Chrysopa 

rufipes, Megaselia 

rufocoxalis, Rogas 

rufopicta, Winthemia 

rugosa, Lysiphlebia 

rugosa, Sphegigaster 

rugosa, Trigonogastra 

ruralis, Voria 

rutila, Agonoscelis 

rutovinctus, Cryptus 


saccharicolus, Blepyrus 

salicaphis, Adialytus 

salicaphis, Lysiphlebus 

salicis, Aphidius 

saltator, Conocephalus 

samoana, Enicospilus 

sanctus, Sympherobius 

sandanis, Sympiesis 

sanguinea, Cycloneda 

Sarcodexia innota (Walker) Dip.: Sarcophagidae 202 

Sarcodexia sternodontis Townsend Dip.: Sarcophagidae 202, 327


Sarcophaga sp. Dip.: Sarcophagidae 

sardus, Horismenus 

sativae, Liriomyza 

saundersii, Madremyia 

sawadai, Anagyrus 

scapuliferus, Scymnus 

328 

Scarites Col.: Carabidae 

scelestes, Brinckochrysa 

scelestes, Chrysopa 

schayeri, Calosoma 

schellenbergii, Oechalia 

schimitscheki, Lysaphidus 

90, 102 

Schizaphis graminum (Rondani) Hem.: Aphididae 5 

Schizobremia formosana Felt Dip.: Cecidomyiidae 147, 154 

Schizocerophaga leibyi Townsend Dip.: Tachinidae 

schwarzi, Euryrhopalus 

328 

Scleroderma Hym.: Bethylidae 180 

Scleroderma cadaverica Benoit Hym.: Bethylidae 168, 180 

Scotinophara lurida (Burmeister) Hem.: Pentatomidae 224 

Scotolinx phyllocnistis, see Cirrospilus phyllocnistis 267 

Scotolinx quadristriata, see Cirrospilus quadristriata 

scutellaris, Ischiodon 

scutellaris, Lipolexis 

scuticarinatus, Trissolcus 

267 

Scymnus Col.: Coccinellidae 43, 66, 146, 150, 152, 293, 305 

Scymnus apetzi Mulsant Col.: Coccinellidae 292, 306 

Scymnus apiciflavus (Motschulsky) Col.: Coccinellidae 146, 292 

Scymnus bilucenarius (Mulsant) Col.: Coccinellidae 145, 146, 150, 152 

Scymnus biguttatus Mulsant Col.: Coccinellidae 292, 306 

Scymnus binaevatus Mulsant Col.: Coccinellidae 293, 296, 310 

Scymnus bipunctatus, see Nephus bipunctatus 292, 296, 310 

Scymnus constrictus Mulsant Col.: Coccinellidae 68 

Scymnus hoffmanni Weise Col.: Coccinellidae 62, 63, 66 

Scymnus includens (Kirsch) Col.: Coccinellidae 292, 314 

Scymnus louisianae Chapin Col.: Coccinellidae 69 

Scymnus margipallens Mulsant Col.: Coccinellidae 150, 154 

Scymnus mauritiusi Korsch. Col.: Coccinellidae 146, 153 

Scymnus pictus Gorham Col.: Coccinellidae 150


Scymnus quadrivittatus Mulsant Col.: Coccinellidae 150 

Scymnus reunioni, see Nephus reunioni 292, 302, 304, 310 

Scymnus roepkei De Fluiter Col.: Coccinellidae 292 

Scymnus scapuliferus Mulsant Col.: Coccinellidae 43 

Scymnus sordidus Horn Col.: Coccinellidae 152, 292, 300, 310 

Scymnus subvillosus (Goeze) Col.: Coccinellidae 292, 306 

Scymnus trepidulus Weise Col.: Coccinellidae 43 

Scymnus uncinatus Sicard Col.: Coccinellidae 145, 150 

semialbicornis, Hemiptarsenus 

Semielacher Hym.: Eulophidae 269, 276 

Semielacher petiolatus (Girault) Hym.: Eulophidae 257, 264, 269, 281, 284, 285 

semiflavus, Aphelinus 

semifumatum, Trichogramma 

Senometopia, see Carcelia 111 

Senotainia Dip.: Sarcophagidae 327 

septempunctata, Chrysopa 

septempunctata, Coccinella 

Sericophoromyia marshalli Villeneuve Dip.: Tachinidae 23 

serratus, Paragus 

serripes, Riptortus 

sexdentatus, Pediobius 

sexmaculata, Cheilomenes 

sexmaculatus, Menochilus 

seychellensis, Telenomus 

shaanxiensis, Lysiphlebus 

shakespearei, Merismorella 

shakespearei, Syntomopus 

side, Spilochalcis 

Sigalphus nigripes He & Chen Hym.: Braconidae 24 

signata, Chrysopa 

Signiphora Hym.: Signiphoridae 123, 129 

silesiaca, Leucopis 

silvestri, Hyperaspis 

similis, Aphidius 

simplex, Chilo 

simplex, Cuspicona 

Sinea complexa Caudell Hem.: Reduviidae 331 

Sinea confusa Caudell Hem.: Reduviidae 331


Sinea diadema (Fabricius) Hem.: Reduviidae 331 

sinensis, Trioxys 

sinica, Chrysopa 

sinica, Chrysoperla 

sinoferus, Nabis 

Siphona Dip.: Tachinidae 328 

Siphona plusiae Coquillett Dip.: Tachinidae 328 

sipius, Trissolcus 

Sitotroga cerealella (Olivier) Lep.: Gelechiidae 344 

smithi, Coccodiplosis 

Snellenius manilae (Ashmead) Hym.: Braconidae 323 

sojae, Melanagromyza 

solani, Phenacoccus 

Solenopsis Hym.: Formicidae 152, 153, 275 

Solenopsis geminata (Fabricius) Hym.: Formicidae 144, 153, 154, 223 

solitarius, Pachyneuron 

solitus, Telenomus 

solocis, Trissolcus 

sonchi, Aphidius 

sonorensis, Campoletis 

sorbillans, Exorista 

sordidus, Cosmopolites 

sordidus, Scymnus 

Spalgis epius (Westwood) Lep.: Lycaenidae 294, 302, 314 

sparsior, Hesperus 

spencerella, Ophiomyia 

Sphaerophoria cylindrica (Say) Dip.: Syrphidae 332 

Sphaerophoria menthastri (Linnaeus) Dip.: Syrphidae 332 

Sphaerophoria robusta (Curran) Dip.: Syrphidae 332 

Sphegigaster Hym.: Pteromalidae 244, 247 

Sphegigaster agromyzae, see Sphegigaster voltairei 254 

Sphegigaster brunneicornis (Ferri re) Hym.: Pteromalidae 243, 245, 246, 254 

Sphegigaster hamygurivara Hym.: Pteromalidae 243 

Sphegigaster rugosa (Waterston) Hym.: Pteromalidae 243, 246, 254 

Sphegigaster stella (Girault) Hym.: Pteromalidae 244, 247, 254 

Sphegigaster stepicola Bou‹ek Hym.: Pteromalidae 244, 245, 254 

Sphegigaster voltairei (Girault) Hym.: Pteromalidae 244, 245, 247, 254 

Spilochalcis flavopicta Cresson Hym.: Chalcididae 323


Spilochalcis nr. mariae (Riley) Hym.: Chalcididae 323 

Spilochalcis side Walker Hym.: Chalcididae 323 

spinator, Pristomerus 

spiraecola, Aphis 

splendens, Chrysoplatycerus 

Spodoptera exigua HŸbner Lep.: Noctuidae 16, 30, 194, 345 

Spodoptera littoralis (Boisduval) Lep.: Noctuidae 30 

stella, Paratrigonogastra 

stella, Sphegigaster 

Stenichneumon culpator cincticornis (Cresson) Hym.: Ichneumonidae 326, 348 

Stenomesius japonicus (Ashmead) Hym.: Eulophidae 269 

Stephanoderes hampei, see Hypothenemus hampei 158 

stephanoderis, Cephalonomia 

stepicola, Sphegigaster 

stercorator orgyiae, Iseropus 

sternodontis, Sarcodexia 

Sticholotis quatrosignata Weise Col.: Coccinellidae 146 

Stictopisthus africanus, see Mesochorus africanus 28 

Stiretrus anchorago (Fabricius) Col.: Pentatomidae 331 

striaditera, Hololepta 

striata, Trathala 

striaticeps, Psix 

striatipes, Sympiesis 

Sturmia atropivora, see Zygobothria atropivora 12 

Sturmia auratocauda, see Cadurcia auratocauda 22 

Sturmia bimaculata, see Palexorista inconspicua 23 

Sturmia dilabida Villeneuve Dip.: Tachinidae 12, 16 

Sturmia macrophallus, see Zygobothria ciliata 12 

Sturmia parachrysops Bezzi Dip.: Tachinidae 190 

subaenea, Oxyharma 

subaenea, Pterosema 

subaeneus, Thysanus 

subdepressum, Dactylosternum 

subimpictus, Phonoctonus 

sublimbalis, Deanolis 

submetallicus, Ooencyrtus 

subquadratum, Dactylosternum 

subvillosus, Scymnus


sulphurea, Cheilomenes 

suppressalis, Chilo 

suturalis, Brumus 

Sycanus indagator StŒl Col.: Reduviidae 

syleptae, Apanteles 

syleptae, Eurytoma 

331 

Syllepte derogata Fabricius Lep.: Pyralidae 28, 194 

Sympherobius barberi (Banks) Neu.: Chrysopidae 291, 295, 298, 305 

Sympherobius sanctus Tjeder Neu.: Chrysopidae 291, 303 

Sympiesis Hym.: Eulophidae 264, 269, 276, 279 

Sympiesis gregori Hym.: Eulophidae 269, 276 

Sympiesis striatipes (Ashmead) Hym.: Eulophidae 270, 274, 275, 277 

Sympiesomorpha mikan, see Stenomesius japonicus 269 

Synopeas rhanis (Walker) Hym.: Platygasteridae 80 

Syntomopus Hym.: Pteromalidae 244 

Syntomopus shakespearei (Girault) Hym.: Pteromalidae 244, 254 

Syrphophagus africanus Gahan Hym.: Encyrtidae 68 

Syrphophagus aphidivora (Mayr) Hym.: Encyrtidae 42, 68, 72 

Syrphophagus taiwanus Hayat and Lin Hym.: Encyrtidae 123, 125 

Syrphus Dip.: Syrphidae 43, 67, 69, 293, 300 

Syrphus balteatus, see Episyrphus balteatus 43 

Syrphus confrater, see Eupeodes confrater 43 

tachinomoides, Euphorocera 

taiwanus, Syrphophagus 

Tamarixia Hym.: Eulophidae 128, 133 

Tamarixia dryi (Waterston) Hym.: Eulophidae 129, 130 

Tamarixia leucaenae Bou‹ek Hym.: Eulophidae 134 

Tamarixia radiata (Waterston) Hym.: Eulophidae 113, 120, 121, 123Ð134 

Tapinoma Hym.: Formicidae 275 

Technomyrmex detorquens Walker Hym.: Formicidae 153 

Telenomus Hym.: Scelionidae 12, 14, 15, 201, 205, 213, 218, 220, 221, 227, 233, 326 

Telenomus acrobates Giard. Hym.: Scelionidae 306 

Telenomus chloropus (Thomson) Hym.: Scelionidae 

227, 229, 234 

200, 205, 211Ð213, 217, 221, 224, 

Telenomus comperei Crawford Hym.: Scelionidae 205, 226 

Telenomus cristatus Johnson Hym.: Scelionidae 205 

Telenomus cyrus Nixon Hym.: Scelionidae 206, 230


Telenomus gifuensis Ashmead Hym.: Scelionidae 206, 212, 221, 224 

Telenomus mormideae Costa Lima Hym.: Scelionidae 206, 220, 221 

Telenomus nakagawai, see Telenomus chloropus 211, 224, 229 

Telenomus pacificus (Gahan) Hym.: Scelionidae 205, 226 

Telenomus podisi (Ashmead) Hym.: Scelionidae 205, 221 

Telenomus seychellensis Kieffer Hym.: Scelionidae 205, 220 

Telenomus solitus Johnson Hym.: Scelionidae 326 

Teleopterus Hym.: Eulophidae 270, 272, 288 

Teleopterus delucchii Bou‹ek Hym.: Eulophidae 270, 278 

tenellus, Gelis 

testaceipes, Lysiphlebus 

testaceus, Pristomerus 

testulalis, Maruca 

tetracanthus, Zelus 

Tetramorium bicarinatum (Nylander) Hym.: Formicidae 85, 92, 100, 101 

Tetramorium guineense, see Tetramorium bicarinatum 92, 100 

Tetrastichus Hym.: Eulophidae 120, 123, 125, 129, 131, 133, 241, 246, 270, 273, 274, 

276, 277, 285 

Tetrastichus ayyari, see Tetrastichus howardi 24 

Tetrastichus howardi (Olliff) Hym.: Eulophidae 24, 29 

Tetrastichus phyllocnistoides, see Citrostichus phyllocnistoides 268 

Tetrastichus radiatus, see Tamarixia radiata 121 

texanus, Chelonus 

thalense, Trichogramma 

theclarum, Aplomya 

theobromae, Aenasius 

theristis, Pammene 

Thopeutis intacta Snellen Lep.: Pyralidae 284 

thyantae, Trissolcus 

Thyreocephalus interocularis Eppelsheim Col.: Staphylinidae 85, 92, 93, 103 

Thysanus niger Ashmead Hym.: Encyrtidae 148 

Thysanus subaeneus Forst. Hym.: Encyrtidae 306 

tibialis, Brachymeria 

tibialis, Microcharops 

tibialis, Paragus 

Timberlakia gilva Prinsloo Hym.: Encyrtidae 295, 298 

timidus, Micromus 

tischeriae, Elasmus


Toxares macrosiphophagum Shuja Uddin Hym.: Aphidiidae 56 

Toxares zakai Shuja Uddin Hym.: Aphidiidae 41 

Toxoptera aurantii (Boyer de Fonscolombe) Hem.: Aphididae 

transcaspicus, Aphidius 

transvena, Encarsia 

transversalis, Coccinella 

transversoguttata, Coccinella 

64, 75, 79 

Trathala flavoorbitalis (Cameron) Hym.: Ichneumonidae 185, 191Ð193, 195 

Trathala striata Cameron Hym.: Ichneumonidae 

trepidulus, Scymnus 

191 

Trichogramma Hym.: Trichogrammatidae 13, 14, 16, 21, 25, 30, 195, 317, 321, 327, 341 

Trichogramma achaeae Nagaraja & Nagarkatti Hym.: Trichogrammatidae 13Ð16 

Trichogramma agriae Nagaraja Hym.: Trichogrammatidae 13Ð16 

Trichogramma australicum Girault Hym.: Trichogrammatidae 13Ð16, 326 

Trichogramma brevicapillum Pinto & Platner Hym.: Trichogrammatidae 326 

Trichogramma chilonis Ishii Hym.: Trichogrammatidae 13, 14, 16, 25, 29, 30, 111 

Trichogramma chilotraeae Nagaraja and Nagarkatti Hym.: Trichogrammatidae 11, 326 

Trichogramma confusum Viggiani Hym.: Trichogrammatidae 13, 15 

Trichogramma deion Pinto & Oatman Hym.: Trichogrammatidae 326 

Trichogramma dendrolimi Matsumura Hym.: Trichogrammatidae 21, 25, 29 

Trichogramma evanescens Westwood Hym.: Trichogrammatidae 326, 344 

Trichogramma exiguum Pinto & Platner Hym.: Trichogrammatidae 326 

Trichogramma japonicum Ashmead Hym.: Trichogrammatidae 25, 30, 326 

Trichogramma minutum Riley Hym.: Trichogrammatidae 

344 

13, 15, 25, 27, 29, 326, 339, 

Trichogramma platneri Nagarkatti Hym.: Trichogrammatidae 326, 344, 346 

Trichogramma pretiosum Riley Hym.: Trichogrammatidae 27, 327, 337, 338, 344, 346 

Trichogramma semifumatum (Perkins) Hym.: Trichogrammatidae 327, 340 

Trichogramma thalense Pinto & Oatman Hym.: Trichogrammatidae 327 

Trichoplusia ni HŸbner Lep.: Noctuidae 1, 2, 317Ð347 

Trichopoda Dip.: Tachinidae 203, 208, 232 

Trichopoda giacomellii (Blanchard) Dip.: Tachinidae 

231, 232, 234 

202, 211, 218, 220, 221, 225, 226, 

Trichopoda gustavoi, see Trichopoda giacomellii 202 

Trichopoda lanipes (Fabricius) Dip.: Tachinidae 202, 232 

Trichopoda nigrifrontalis, see Trichopoda giacomelli 202 

Trichopoda pennipes (Fabricius) Dip.: Tachinidae 

221, 224Ð226, 230, 231, 233 

197, 201, 208, 209, 212Ð215, 217,


Trichopoda pilipes (Fabricius) Dip.: Tachinidae 

233 

trifasciatus, Closterocerus 

trifolii, Liriomyza 

197, 203, 211Ð214, 217, 221, 225, 231, 

Trigonogastra agromyzae, see Sphegigaster voltairei 245, 254 

Trigonogastra rugosa, see Sphegigaster rugosa 

trilongifasciatus, Leptomastix 

trinidadensis, Ooencyrtus 

254 

Triommata coccidivora (Felt) Dip.: Cecidomyidae 293, 302 

Trioxys Hym.: Aphidiidae 42, 57 

Trioxys acalephae (Marshall) Hym.: Aphidiidae 41, 56 

Trioxys angelicae (Haliday) Hym.: Aphidiidae 41, 56, 65, 71, 78 

Trioxys asiaticus Telenga Hym.: Aphidiidae 41, 56 

Trioxys auctus (Haliday) Hym.: Aphidiidae 41, 56 

Trioxys basicurvus Shuja Uddin Hym.: Aphidiidae 56, 64 

Trioxys centaureae (Haliday) Hym.: Aphidiidae 41 

Trioxys cirsii (Curtis) Hym.: Aphidiidae 41 

Trioxys communis Gahan Hym.: Aphidiidae 56, 63, 66Ð68, 78, 82, 83 

Trioxys complanatus Quilis Hym.: Aphidiidae 42, 56 

Trioxys equatus Samanta, Tamili and Raychaudhuri Hym.: Aphidiidae 56 

Trioxys hokkaidensis Takada Hym.: Aphidiidae 42 

Trioxys indicus Subba Rao & Sharma Hym.: Aphidiidae 42, 57, 61Ð65, 78, 79, 82, 83 

Trioxys nearctaphidis (Mackauer) Hym.: Aphidiidae 42 

Trioxys pallidus Haliday Hym.: Aphidiidae 57 

Trioxys rietscheli Mackauer Hym.: Aphidiidae 57, 68 

Trioxys rubicola Shuja Uddin Hym.: Aphidiidae 64 

Trioxys sinensis Mackauer Hym.: Aphidiidae 42, 57, 67 

Trioza citroimpura Yang and Li Hem.: Psyllidae 116 

Trioza eastopi, see Trioza litseae 129 

Trioza erytreae (Del Guerico) Hem.: Psyllidae 119, 120, 129, 130 

Trioza litseae Bordaga Hem.: Psyllidae 129 

Trissolcus Hym.: Scelionidae 201, 206, 211, 221, 226, 227, 233 

Trissolcus aloysiisabaudiae (Fouts) Hym.: Scelionidae 206, 220 

Trissolcus basalis (Wollaston) Hym.: Scelionidae 197, 201, 206, 209, 211Ð228, 230 

Trissolcus brochymenae (Ashmead) Hym.: Scelionidae 206 

Trissolcus crypticus Clarke Hym.: Scelionidae 207, 210, 211 

Trissolcus euschisti Ashmead Hym.: Scelionidae 207 

Trissolcus hullensis (Harrington) Hym.: Scelionidae 207


Trissolcus lepelleyi (Nixon) Hym.: Scelionidae 207, 220 

Trissolcus lodosi (Szab—) Hym.: Scelionidae 207 

Trissolcus maro Nixon Hym.: Scelionidae 207, 220 

Trissolcus mitsukurii (Ashmead) Hym.: Scelionidae 206, 211Ð215, 217, 221, 224, 229 

Trissolcus oenone (Dodd) Hym.: Scelionidae 206 

Trissolcus ogyges (Dodd) Hym.: Scelionidae 206 

Trissolcus rudus Le Hym.: Scelionidae 206 

Trissolcus scuticarinatus (Costa Lima) Hym.: Scelionidae 206 

Trissolcus sipius (Nixon) Hym.: Scelionidae 206, 220 

Trissolcus solocis Johnson Hym.: Scelionidae 206 

Trissolcus thyantae Ashmead Hym.: Scelionidae 206 

Trissolcus urichi Hym.: Scelionidae 206 

Trissolcus utahensis Hym.: Scelionidae 

tristicolor, Orius 

tristis, Anasa 

206 

Trogus exaltatorius Panzer Hym.: Ichneumonidae 12 

Tropidophryne melvillei Compere Hym.: Encyrtidae 

truncatellum, Copidosoma 

309 

ultimus, Gambrus 

uncinatus, Scymnus 

undecimpunctata, Coccinella 

unicolor, Leptochirus 

unicolor, Priochirus 

urichi, Trissolcus 

urozonus, Eupelmus 

urticae, Aphidius 

utahensis, Trissolcus 

uzbekistanicus, Aphidius 

v-nigrum, Olla 

varicornis, Hemiptarsenus 

variegata, Adonia 

variegata, Hippodamia 

variegatus, Cirrospilus 

varipes, Aphelinus 

venator, Lissauchenius 

ventralis, Rhizobius


Vespula pensylvanica (Saussure) Hym.: Vespidae 333 

vicina, Cheilomenes 

Vincentodiplosis pseudococci (Felt) Dip.: Cecidomyiidae 145, 151, 156 

viridicoxa, Callitula 

viridicoxa, Eurydinotellus 

viridula, Nezara 

Visnuella brevipetiolatu, see Zaommomentedon brevipetiolatus 270, 275 

vitis, Pseudococcus 

vitodurense, Pachyneuron 

vitrata, Maruca 

vitripennis, Glyptapanteles 

vittatus, Cirrospilus 

vittatus, Collops 

vittegera, Paranaemia 

voltairei, Sphegigaster 

volucre, Praon 

Voria edentata (Baranof) Dip.: Tachinidae 328 

Voria ruralis (FallŽn) Dip.: Tachinidae 317, 328, 336Ð341, 345Ð347 

vulgaris, Phryxe 

Vulgichneumon brevicinctor (Say) Hym.: Ichneumonidae 326, 348 

walkeri, Chartocerus 

wallacii, Cryptolaemus 

Wasmannia auropunctata (Roger) Hym.: Formicidae 174 

Winthemia Dip.: Tachinidae 329, 337 

Winthemia dasyops (Wiedemann) Dip.: Tachinidae 21, 23, 28 

Winthemia montana Rein. Dip.: Tachinidae 328 

Winthemia pinguis Dip.: Tachinidae 337 

Winthemia pyrrhopyga (Wiedemann) Dip.: Tachinidae 337 

Winthemia quadripustulata (Fabricius) Dip.: Tachinidae 328 

Winthemia rufopicta (Bigot) Dip.: Tachinidae 329 

Xanthodes graellsii (Feisthamel) Lep.: Noctuidae 30 

Xanthogramma aegyptium, see Ischiodon aegyptius 43 

Xanthopimpla punctata (Fabricius) Hym.: Ichneumonidae 25, 191, 192 

Xenoencyrtus Hym.: Encyrtidae 204 

Xenoencyrtus hemipterus (Girault) Hym.: Encyrtidae 204, 211, 213, 214, 218, 221 

Xenoencyrtus niger, see Xenoencyrtus hemipterus 204, 211, 213, 221


Xenoencyrtus rubricatus Riek Hym.: Encyrtidae 204 

xylostella, Plutella 

yakutatensis, Cotesia 

yasudi, Callitula 

Zacharops narangae Cushman Hym.: Braconidae 25 

Zagrammosoma Hym.: Braconidae 278 

Zagrammosoma multilineatum (Ashmead) Hym.: Eulophidae 

zakai, Toxares 

270, 273, 278 

Zaommomentedon Hym.: Eulophidae 270, 276 

Zaommomentedon brevipetiolatus Kamijo Hym.: Eulophidae 270, 275, 277, 281, 285 

Zaplatycerus fullawayi Timberlake Hym.: Encyrtidae 

zehntneri, Elasmus 

149 

Zelus bilobus Say Hem.: Reduviidae 331 

Zelus exsaguis StŒl Hem.: Reduviidae 331 

Zelus renardii Kalenati Hem.: Reduviidae 331 

Zelus tetracanthus StŒl Hem.: Reduviidae 331 

Zenilla blanda blanda (Osten Sacken) Dip.: Tachinidae 329 

Zenillia cosmophilae, see Carcelia cosmophilae 22 

Zenillia noctuae, see Carcelia illota 

zizyphi, Aphis 

20 

Zygobothria atropivora (Robineau-Desvoidy) Dip.: Tachinidae 12, 16 

Zygobothria ciliata (Wulp) Dip.: Tachinidae 12, 15, 23

7 General Index 

abelmoschus, Hibiscus 

aconitifolia, Vigna 

Acridotheres tristis 26 

Aegle marmelos 262 

alatae 34, 35, 46, 70 

alfalfa looper, see Autographa californica 342 

Allothrombium pulvinum 58, 63 

Alseodaphne semicarpifolia 262 

Althaea rosea 19 

Amaranthus 19 

Ananas comosus 142 

anise, see Clausena anisumolens 

anisopliae, Metarhizium 

anisumolens, Clausena 

annuus, Helianthus 

antiquorum, Colocasia 

117 

Apocynum venotum 29 

apterae 34, 46, 47 

Arabian coffee, see Coffea arabica 159, 165, 181 

arabica, Coffea 181 

Arbutilon 19 

areca palm 144 

Argentine ant, see Iridomyrmex humilis 144 

Arthrobotrys 59 

artificial diet 11, 35, 170, 177, 187, 320 

arvensis, Convolvulus 

Asian citrus psyllid, see Diaphorina citri 

asiatica, Toddalia 

114 

asparagus 320 

Aspergillus flavus 

astrigata, Pardosa 

138, 298, 301, 334 

Atalantia 118 

atropurpureum, Macroptilium 

atropurpureus, Phaseolus 

aurantiifolia, Citrus 

aurea, Vigna 

531


aureus, Phaseolus 

australisiaca, Microcitrus 

avocado 108 

Bacillus thuringiensis 21, 27, 320, 334 

Bacillus thuringiensis wuhanensis 

bacteriophora, Heterorhabditis 

26 

baculovirus 191, 342 

banana, see Musa sapientum 68, 85--104, 144, 145, 154, 163, 289 

banana aphid, see Pentalonia nigronervosa 67, 68, 74 

banana weevil borer, see Cosmopolites sordidus 85, 86 

barley 5 

bassiana, Beauveria 

batatas, Ipomoea 

bats 321 

bean, see Phaseolus vulgaris 

238, 251, 320 

35, 47, 63, 161, 199, 210, 219, 236, 

bean fly, see Ophiomyia phaseoli 235, 236 

bean stem borer, see Melanagromyza sojae 

Beauveria 128, 167, 199 

Beauveria bassiana 59, 69, 93, 122, 128, 157, 166, 169, 170, 174-- 

176, 182, 183, 334 

beet armyworm, see Spodoptera exigua 

bele, see Hibiscus manihot 

Bergera koenigii 116, 117 

19 

bigheaded ant, see Pheidole megacephalaI 144 

bindweed, see Convulvulus arvensis 11 

black berry nightshade, see Solanum nigrum 187 

black citrus aphid, see Toxoptera aurantii 75 

black gram, see Vigna mungo 237 

black legume aphid, see Aphis craccivora 34 

bluish dogbane, see Apocynum venotum 29 

Botrytis stephanoderis see Beauveria bassiana 169 

bottle gourd 78 

Bouea burmanica 105, 108 

brinjal, see Solanum melongena 185 

brinjal fruit borer, see Leucinodes orbonalis 186 

brinjal stem borer, see Euzophera ferticella 194 

broccoli 321

unneum, Metarrhizium 

burmanica, Bouea 

cabbage 317, 320, 321, 337 

cabbage aphid, see Brevicoryne brassicae 48 

cabbage looper, see Trichoplusia ni 317, 318 

cabbage white butterfly, see Pieris rapae 

caerulea, Geoplana 

caerulea, Kontikia 

337 

Caesalpinia 163 

cairica, Ipomoea 

cajan, Cajanus 

Cajanus cajan 

calcarata, Vigna 

237 

caloxylon, Merrillia 

Canavalia ensiformis 

canephora, Coffea 

cannabinus, Hibiscus 

238 

canola 320 

cantaloupe 320 

cape gooseberry 187 

Capsella 35 

capsicum 187, 200, 320 

carcocapsae, Steinernema 

carrot 320 

cassava 289 

castor, see Ricinus communis 232 

Catharanthus roseus 116 

catimor coffee 176 

cauliflower 320 

celery 320, 341 

Centrosema 163 

Cephalosporium lecanii 59, 69, 122, 128 

Chenopodium 338 

chick pea 236 

chico 108 

chinense, Clerodendrum 

Chinese cabbage 63 

Chiracanthium inclusum 278 

General Index 533


chlordimeform 27 

chrysanthemum 5, 65 

Ciconia nigra 16 

Cinnamomum zeylanica 

cinnamomum, Jasminum 

citri, Xanthomonas 

262 

citricola scale, see Coccus pseudomagnoliarum 299 

citron, see Citrus medica 117 

Citrullus lanatus 19 

Citrus 35, 47, 64--66, 77, 113--134, 257--316 

Citrus aurantifolia 117, 262 

Citrus hystrix 117 

Citrus limon 117, 262 

Citrus madurensis 117 

Citrus maxima 117 

Citrus maxima var. racemosa 117 

Citrus medica 117, 262 

Citrus reticulata 115, 117 

Citrus sinensis 117 

citrus canker fungus, see Xanthomonas citri 263 

citrus greening 113, 114, 116, 119, 120, 128--130 

citrus leaf mottle 128, 130 

citrus leafminer, see Phyllocnistis citrella 257, 258 

citrus mealybug, see Planococcus citri 287, 288 

citrus psyllid, see Diaphorina citri 114 

citrus tristeza virus 36, 47 

citrus vein phloem degeneration 128 

Cladosporium oxysporum 298, 305 

Clausena anisumolens 116, 117 

Clausena excavata 117 

Clausena indica 118 

Clausena lansium 117 

Clerodendrum chinense 14, 15 

Clubiona 278 

cocoa 35, 47, 289 

coconut bug, see Amblypelta cocophaga 226 

coconut spathe bug, see Axiagastus campbelli 

coerulia, Tweedia 

226

General Index 535 

Coffea 163, 288 

Coffea arabica 158, 159, 165, 176, 181 

Coffea canephora 159, 165, 181 

coffeanum, Colleotrichium 

coffee 35, 47, 144, 157--183, 289, 290, 302--304 

coffee berry borer, see Hypothenemus hampei 158 

coffee berry disease, see Colletotrichum coffeanum 170 

coffee leaf rust, see Hemileia vastatrix 170, 176 

Colleotrichium coffeanum 170 

Colocasia antiquorum 14, 15 

Colocasia esculenta 

communis, Ricinus 

comosus, Ananas 

71 

Convolvulus arvensis 11 

cotton 17, 19, 21, 27--30, 46, 47, 60, 62, 63, 65, 68, 69, 78, 137, 161, 

200, 220, 320, 335, 338, 340--345, 347 

cotton aphid, see Aphis gossypii 46 

cotton semi looper, see Anomis flava 18 

cotton stainer, see Dysdercus cingulatus 135, 136 

cowpea, see Vigna unguiculata or Vigna sinensis 

219, 222, 227, 236, 238, 239 

19, 35, 36, 200, 

cowpea aphid, see Aphis craccivora 34 

crazy ant, see Paratrechina longicornis 145 

creeping sensitive plant, see Mimosa invisa 

croceus, Xysticus 

134 

Crotalaria 163, 251--254 

Crotalaria juncea 238 

Crotalaria laburnifolia 238, 245 

Crotalaria mucronata 238 

cucumber 47, 64, 66, 67, 73, 83, 187, 320 

curry bush, see Bergera koenigii 117 

custard apple 299, 300 

Cyclosa insulana 65 

cyfluthrin 112


date 289 

DD136 Steinernema carcocapsae 138 

Deccan hemp, see Hibiscus cannabinus 19 

deltamethrin 112 

Dendroica palmarum 333 

desi cotton 19 

Dialium lacourtianum 163 

diamondback moth, see Plutella xylostella 

diazoma, Thelohania 

337 

Dioscorea 164 

dissecta, Merremia 

Dolichos lablab, see Lablab niger 238 

egg plant, see Solanum melongena 47, 64, 65, 78, 83, 185--195 

egg plant fruit and shoot borer, see Leucinodes orbonalis 186 

elephant lemon, see Citrus medica 262 

endosulfan 180, 182 

ensiformis, Canavalia 

Entomophthora 13, 59, 335 

Entomophthora exitialis 59 

Entomophthora fresenii 298 

Entomophthora fumosa 298, 300 

Entomophthora gammae 334 

Entomophthora sp. ÔgrylliÕ type 13 

Entomophthora sphaerosperma 334 

Eremocitrus glauca 262 

Erigonidium graminicolum 

esculenta, Colocasia 

esculentum, Lycopersicon 

esculentus, Hibiscus 

excavata, Clausena 

exitialis, Entomophthora 

exotica, Murraya 

26, 62


faba, Vicia 

feltiae, Steinernema 

fig 288, 289, 306 

fire ant, see Solenopsis geminata 144 

flavus, Aspergillus 

flax 231 

Fortunella 116, 117 

French bean, see Phaseolus vulgaris 237, 245, 249, 253 

fresenii, Entomophthora 

fresenii, Neozygites 

fruit sucking moth 19 

fumosoroseus, Paecilomyces 

Fusarium 298 

Fusarium lateritium 122, 128 

Garcinia mangostana 262 

gardenia 290 

Geoplana caerulea see Kontikia caerulea 100 

glasshouse 35, 59, 66, 69, 71, 73, 77, 288, 290, 305, 315, 316 

glauca, Eremocitrus 

glauca, Nicotiana 

glaucum, Pennisetum 

Gliricidia 63 

Gliricidia maculata 68 

Gliricidia sepium 

glutinesa, Swinglea 

35, 164 

Glycine max 67, 219, 238, 245, 246 

Glycine soja 248 

golden gram, see Vigna aurea 

graminicolum, Erigonidium 

237 

granulosis virus 21, 26 

grape 19, 119, 199, 288--290, 306, 315 

grapefruit 303, 305 

green gram, see Vigna aurea or Vigna radiata 19, 237, 247 

green lacewing, see Chrysoperla carnea 43 

green peach aphid, see Myzus persicae 74 

green semi looper, see Anomis flava 18 

green vegetable bug, see Nezara viridula 198 

groundnut 35, 144, 199, 226 

groundnut aphid, see Aphis craccivora 34


guava 47, 108, 200, 302 

gypsy moth 225 

Habana velox 278 

haricot bean, see Phaseolus vulgaris 

hederifolia, Ipomoea 

237 

Helianthus annuus 219 

Hemileia vastatrix 170 

Hemipterotarseius 137, 138 

Hentzia palmarum 279 

Heterorhabditis 93, 94, 167, 169 

Heterorhabditis bacteriophora 94 

Heterorhabditis zealandica 94 

Hibiscus 60, 64, 76, 83, 137, 163 

Hibiscus abelmoschus 19 

Hibiscus cannabinus 19 

Hibiscus esculentus 19, 67 

Hibiscus manihot 19 

Hibiscus rosa-sinensis 19 

Hibiscus sabadariffa 19 

hirsutum cotton 19 

hollyhock, see Althaea rosea 19 

honey 226, 284, 313, 344 

honeydew 47, 74, 82, 132, 144, 156, 290, 303, 315 

horehound bug, see Agonoscelis rutila 218 

huanglunbin 116 

hyacinth bean, see Lablab niger 

hystrix, Citrus 

238 

Indian mynah 21, 29 

indica, Clausena 

indica, Ipomoea 

indicum, Sesamum 

indicum, Solanum 

insulana, Cyclosa 

invisa, Mimosa 

Ipomoea 15 

Ipomoea batatas 11, 19 

Ipomoea cairica 11

Ipomoea hederifolia 11 

Ipomoea indica 11 

Ipomoea pescapreae 11 

jackfruit 108 

jasmin orange, see Murraya paniculata 116, 117 

Jasminum cinnamomum 262 

Jasminum humile 262 

Jasminum sambac 262 

javanicus, Paecilomyces 

javanicus, Spicaria 

jute, see Hibiscus sabadariffa 19, 230 

kapok, see Hibiscus cannabinus 137 

kenaf 17, 19, 27, 30 

kidney bean, see Phaseolus vulgaris 

koenigii, Bergera 

koenigii, Murraya 

76 

Kontikia caerulea 100 

kumquat, see Fortunella 117 


lablab, see Lablab niger 200, 246 

Lablab niger 238 

laburnifolia, Crotalaria 

lacourtianum, Dialium 

lanatus, Citrullus 

lanceolata, Vepris 

lansium, Clausena 

lateritium, Fusarium 

lathryoides, Macroptilium 

lathyroides, Phaseolus 

leaf mottle 128 

lecanii, Cephalosporium 

lecanii, Verticillium 

Leea 19 

lemon, see Citrus limon 116, 117, 130, 262, 289, 302, 312 

Leonurus sibericus 232 

lettuce 317, 320, 338, 340 

lettuce necrotic yellows 62


Leucaena 163 

Leucaena leucocephala 164 

Ligustrum 163 

likubin 130 

lime, see Citrus aurantifolia 

limon, Citrus 

117, 130, 262, 274, 302, 312 

little red fire ant, see Wasmannia auropunctata 174 

looplure 319 

loquat 164 

Loranthus 262 

lucerne 35, 199, 338, 339, 344 

lunatus, Phaseolus 

lupin, see Lupinus 35 

Lupinus 35 

Lycopersicon esculentum 19, 187 

macadamia 19, 220, 222--224, 289 

Macroptilium atropurpureum 238 

Macroptilium lathryoides 

maculata, Gliricidia 

238 

Madagascar periwinkle, see Catharanthus roseus 

madurensis, Citrus 

116 

maize 14, 137, 161, 199, 320 

Malva 338 

mandarin, see Citrus reticulata 115, 117, 302 

Mangifera 108 

Mangifera indica 105, 106, 108, 109 

Mangifera minor 105, 108 

Mangifera odorata 105, 108 

mango, see Mangifera indica 

mangostana, Garcinia 

47, 105--112, 187, 289, 290 

mangosteen, see Garcinia mangostana 

manihot, Hibiscus 

262 

Manila hemp, see Musa textilis 

marcescens, Serratia 

marmelos, Aegle 

89, 137 

Marpissa tigrina 122, 133 

Mauritius papeda, see Citrus hystrix 117


max, Glycine 35 

maxima var. racemosa, Citrus 

maxima, Citrus 

medic, see Medicago 

medica, Citrus 

Medicago 35 

Melanospora 128 

melon, see Citrullus lanatus 19, 65 

melon aphid, see Aphis gossypii 

melongena, Solanum 

46 

Merremia dissecta 11 

Merrillia caloxylon 117 

Metarhizium 167 

Metarhizium anisopliae 93, 169, 175, 334 

Metarrhizium brunneum 334 

methidathion 302 

methomyl 130 

Microcitrus australisiaca 117 

Mimosa invisa 134 

minor, Mangifera 

Misumenops tricuspidatus 26, 62 

Misumena ratia 333 

Mohinga 101 

morning glory, see Ipomoea hederifolia 11 

moth bean, see Vigna aconitifolia 

mucronata, Crotalaria 

11, 237 

mung bean, see Vigna radiata or Vigna aurea 

mungo, Vigna 

11, 219, 237, 248 

Murraya exotica 118, 262 

Murraya koenigii, see Bergera koenigii 117, 262 

Murraya paniculata 115--117, 127--131, 133 

Musa 86 

Musa sapientum 89 

Musa textilis 89 

muskmallow, see Hibiscus abelmoschus 19 

mustard 338 

myriacanthum, Solanum


Neoaplectana carpocapsae, see Steinernema feltiae 194 

Neozygites fresenii 59, 69 

Nerium 60 

Nicotiana glauca 

niger, Lablab 

nigra, Ciconia 

nigrum, Solanum 

338, 339 

Nomuraea rileyi 169, 334, 335 

Nosema trichoplusiae 334 

octomaculatum, Theridion 

odorata, Mangifera 

oil palm 

okra, see Hibiscus esculentus 17, 19, 35, 47, 65, 137, 227 

okra semi looper, see Anomis flava 18 

oleander aphid, see Aphis nerii 73, 76 

oleander, see Nerium 60, 77 

orange, see Citrus sinensis 117, 199, 302, 304 

oriental stink bug, see Nezara antennata 233 

oviparae 34 

Oxyanthus 163 

Oxyopes salticus 323 

oxysporum, Cladosporium 

Paecilomyces 122, 128 

Paecilomyces fumosoroseus 59, 69 

Paecilomyces javanicus 167, 169 

Paecilomyces tenuipes 

palmarum, Dendroica 

169 

Panagrolaimus 169 

Pandanus 144 

panduratus, Phaseolus 

paniculata, Murraya 

papaya 108 

paprika 35, 47 

paraffin oil 94 

Pardosa 322 

Pardosa astrigata 63


parsley 320 

Passerculus sandwichensis 333 

passionfruit 199, 289, 299, 300 

pea, see Pisum sativum 187, 199, 238, 246, 320 

peanut 161 

pear 19 

pearl millet, see Pennisetum glaucum 137 

pecan 199, 234 

Pennisetum americanum, see Pennisetum glaucum 137 

Pennisetum glaucum 137 

petroleum oil 119, 133, 263, 264, 286 

pescapreae, Ipomoea 

Phaseolus 11, 163, 237 

Phaseolus aureus 67, 247 

Phaseolus lunatus 164 

Phaseolus panduratus 238 

Phaseolus semierectus 238 

Phaseolus vulgaris 76, 219, 236--238, 245 

pheromone 87, 112, 137, 161, 187, 208, 217, 218, 259, 286, 289, 

Phidippus 

319 

333 

Phidippus regius 333 

picorna virus 201 

pigeon pea, see Cajanus cajan 6, 35, 64, 77--79, 236, 237 

pineapple, see Ananas comosus 141--156 

pineapple mealybug wilt 144, 145, 152, 156 

pineapple mealybug, see Dysmicocus brevipes 142 

pineapple scale, see Diaspis bromeliae 153 

Pisum sativum 238 

planarian 100 

plantain 89 

plant resistance 5, 6, 11, 88, 187, 238, 262, 320 

polyacrylamide gel 95 

polyhedrosis virus 21, 26, 317, 321, 334, 337, 338, 340 

pomegranate 288, 306 

Poncirus 116 

Poncirus trifoliata 117 

Pongamia semicarpifolia 262


potato, see Solanum tuberosum 

pseudocacia, Robinia 

47, 65, 187, 199, 289, 320 

prothiophos 94 

Psychotria 164 

pulvinum, Allothrombium 

pummelo, see Citrus maxima var. racemosa 116, 117, 260, 277 

pumpkin, butternut 289 

radiata, Vigna 

rape, see canola 

ratia, Misumena 

320 

red banded borer, see Deanolis sublimbalis 106 

red banded mango caterpillar, see Deanolis albizonalis 106 

red cotton bug, see Dysdercus cingulatus 136 

red seed bug, see Dysdercus cingulatus 

regius, Phidippus 

reticulata, Citrus 

136 

Rhamnus 64, 83 

rhodes grass scale, see Antonina graminis 155 

rice 144, 200, 230, 238 

rice bean, see Vigna calcarata 237 

Ricinus 19 

Ricinus communis 232 

rikettsia 334 

rileyi, Nomuraea 

Robinia pseudacacia 76 

robusta coffee, see Coffea canephora 

rosa-sinensis, Hibiscus 

rosea, Althaea 

159, 165, 181 

roselle, see Hibiscus sabadariffa 19 

roselle cotton 18 

roseus, Catharanthus 

Rubus 163 

Rumex 35 

runner bean, see Phaseolus vulgaris 237


sabadariffa, Hibiscus 

safflower 194 

salticus, Oxyopes 

sambac, Jasminum 

sandwichensis, Passerculus 

santol 108 

sapientum, Musa 

semicarpifolia, Alseodaphne 

semicarpifolia, Pongamia 

semierectus, Phaseolus 

sepium, Gliricidia 

Serratia marcescens 334 

sesame, see Sesamum indicum 200, 219 

Sesamum indicum 219 

Seville orange 286 

shooflower, see Hibiscus rosa-sinensis 

sibericus, Leonurus 

19 

Sida 19 

sineguelas 108 

sinensis, Citrus 

sinensis, Vigna 

sisal 144 

snap bean, see Phaseolus vulgaris 

soja, Glycine 

237 

Solanum 187 

Solanum indicum 187 

Solanum melongena 186, 187 

Solanum myriacanthum 187 

Solanum nigrum 187 

Solanum tuberosum 187 

Solanum xanthocarpum 187 

sooty mould 44, 144, 300 

sordidin 87 

sorghum 5, 137, 199, 218, 230 

southern green stink bug, see Nezara viridula 198 

soybean, see Glycine max or G. soja 67, 144, 177, 199, 200, 201, 

218, 219, 221--224, 226, 227, 229, 231, 232, 234, 236-- 

238, 245--248, 320


soybean stem borer, see Melanagromyza sojae 247 

soybean top borer, see Melanagromyza dolichostigma 247 

soybean, Davis 234 

soybean, PI 717444 200, 234 

sphaerosperma, Entomophthora 

Spicaria javanicus, see Paecilomyces javanicus 

Spicaria see Paecilomyces 

169 

spinach 199, 320 

squash 46, 47, 76 

squash bug, see Anasa tristis 230 

star apple 108 

Steinernema 93, 94 

Steinernema carpocapsae 94, 138, 194, 263 

Steinernema feltiae, see Steinernema carpocapsae 

stephanoderis, Botrytis 

94 

strawberry 69 

subterranean clover 35 

sugarbeet 337 

sugarcane 77, 144, 145, 152 

sunflower, see Helianthus anuus 65, 194, 199, 219 

swallow 166 

sweet potato, see Ipomoea batatas 9, 11, 14, 16, 19, 187 

sweet potato hawk moth, see Agrius convolvuli 10, 14 

sweet potato hornworm, see Agrius convolvuli 10, 14 

sweet potato weevil, see Cosmopolities sordidus 11 

Swinglea glutinesa 118 

tangerine 277 

taro, see Colocasia esculenta 11, 47, 64, 71, 83, 153 

tenuipes, Paecilomyces 

Tephrosia 163 

textilis, Musa 

Thelohania diazoma 334 

Theridion 65 

Theridion octomaculatum 62 

thuringiensis wuhanensis, Bacillus 

thuringiensis, Bacillus 

tigrina, Marpissa


tobacco 199, 289, 320 

Toddalia asiatica 117 

tomato, see Lysopersicum esculentum 19, 187, 195, 199, 200, 210, 

toti virus 

218, 219, 223, 227, 317, 320, 335, 338, 344 

201 

Trachelas volutus 278 

transgenic line 320 

tree tobacco, see Nicotiana glauca 

trichoplusiae, Nosema 

tricuspidatus, Misumenops 

trifoliata, Poncirus 

trifoliata, Triphasia 

338, 339 

Trifolium 35 

Triphasia trifoliata 117 

tristeza virus 34, 47 

tristis, Acridotheres 

tuberosum, Solanum 

Tweedia coerulea 74 

Uloborus 65 

ungiculata, Vigna 

urd bean, see Vigna mungo 11, 237 

Urena 19, 137 

vastatrix, Hemileia 

venotum, Apocynum 

Vepris lanceolata 118 

Verticillium lecanii, see Cephalosporium lecanii 

vexillata, Vigna 

59, 69, 122, 128 

Vicia 35 

Vicia faba 67 

Vigna 237 

Vigna aconitifolia 237 

Vigna aurea 237 

Vigna calcarata 237 

Vigna mungo 11, 14, 237 

Vigna radiata 11, 14, 19, 219, 238 

Vigna sinensis 248


Vigna unguiculata 19, 68, 219, 238, 239, 245 

Vigna vexillata 11 

Vitis 163 

vivapary 34 

vulgaris, Phaseolus 

watermelon 47, 320 

wheat 231 

white-bellied stork, see Ciconia nigra 16 

wild radish 227 

wild mung 11 

woolly apple aphid, see Eriosoma lanigerum 72 

xanthocarpum, Solanum 

Xanthomonas citri 263 

Xysticus croceus 62 

zealandica, Heterorhabditis 

zeylanica, Cinnamomum

Biological Control of Insect Pests: Southeast Asian Prospects - EcoPort

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