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Biological Control of Weeds: 

Southeast Asian Prospects 

D.F. Waterhouse 

(ACIAR Consultant in Plant Protection) 

ACIAR 

(Australian Centre for International Agricultural Research) 

Canberra 

AUSTRALIA 

1994

The Australian Centre for Intemational 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 ACIAR's research objectives. The series is distributed internationally, with an emphasis on the Third 

World. 

O Australian Centre for Intemational Agricultural Research 

GPO Box 157 1, Canberra, ACT 260 1. 

Waterhouse, D.F. 1994. Biological Control of Weeds: Southeast Asian Prospects 

ACIAR Monograph No. 26, vi + 302pp., 27 figs., 28 maps 

ISBN . 186320099 1 

Typset in 11/13 Times using a Macintosh IIvx running Quark XPress by K & B Publications 

Printed by Bmwn Prior Anderson Pty. Ltd.

Foreword 

Contents 

Abstract 

Estimation of biological control prospects 

Introduction 

Target weeds 

Ageratum conyzoides 

Amaranthus spinosus 

Bidens pilosa 

Chromolaena odorata 

Commelina benghalensis 

Echinochloa crus-galli 

Eichhornia crassipes 

Eleusine indica 

Euphorbia heterophylla 

Euphorbia hirta 

Fimbristylis miliacea 

Marsilea minuta 

Melastoma rnalabathricum 

Mikania micrantha 

Mimosa invisa 

Mimosa pigra 

Mimosa pudica 

Monochoria vaginalis 

Nephrolepis biserrata 

Panicum repens 

Paspalurn conjugatum 

PassiJlora foetida 

Pennisetum polystachion 

Pistia stratiotes 

Portulaca oleracea 

Rottboellia cochinchinensis 

Sphenoclea zeylanica 

References 

Index of scientific names of insects 

General index

Foreword 

From its very beginning in 1982 ACIAR has been a strong supporter of biological control 

as a sustainable and environmentally friendly alternative to the steadily growing use of 

pesticides. This alternative has achieved great success in regions of the world 

(e.g. Australia, New Zealand, Oceania, California) where many of the major insect pests 

and weeds have been introduced from outside the region. Although a smaller proportion 

of the major weeds in Southeast Asia are introduced than in many other regions, a recent 

survey commissioned by ACIAR (Waterhouse 1993a) identified 28 major weeds that 

merited evaluation as possible targets for biological control. Even if only half of these 

weeds proved to be attractive targets, this number would require several decades of 

research, major resources in personnel and equipment and strong support within the 

region. 

The aim of the present volume is to summarise for the major exotic weeds of agri- 

culture in Southeast Asia what is known about their natural enemies and the prospects for 

classical biological control. The book is intended to serve two purposes. Firstly, to facili- 

tate, for the countries of the region, the selection of promising, individual or collabora- 

tive, priority weed targets. Secondly, to provide donor agencies with an overall perspec- 

tive of the region's major exotic weed problems and prospects for their amelioration; and 

thus to aid in the selection of projects for support that are best suited to their terms of ref- 

erence. 

It is hoped that it may be possible in the near future to produce a companion volume 

dealing with major arthropod pests exotic to Southeast Asia. 

G.H.L. Rothschild 

Director 

Australian Centre for International 

Agricultural Research, Canberra

Abstract 

Biological control programs have already been mounted in some region of the world 

against 6 of the 28 major weeds that are exotic to Southeast Asia. Substantial or partial 

success has been achieved in one or more countries for all of these except Mikania 

micrantha, which is still under investigation. A substantial amount of information on 

their natural enemies in the region where the weeds evolved is available on all 6. This is 

in stark contrast with the situation for most of the remaining 22 weed species. Indeed, for 

more than half of these, so little relevant information is available that it is not possible to 

evaluate the chances of mounting a successful program. For this group of weeds the first 

step would be a survey in the centre of origin of the weed. It is probable that surveys 

could be mounted simultaneously of several candidate weeds in the same region of the 

world (e.g. Central America or Tropical Africa). The very minimum period for a prelimi- 

nary survey would be several weeks in both spring and late summer. When the organisms 

collected had been identified by taxonomists a decision would be facilitated on possible 

follow-up surveys. 

On the basis of available information there are good to excellent prospects for 

reducing, in at least some parts of the region, the weediness of the following: 



Mimosa invisa 

Mimosa pigra 



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

enemies for the following: 



Bidens pilosa 


Melastoma malabathricum 


There is insufficient information yet available on the remaining 15 weeds to attempt 

to evaluate their prospects for classical biological control.

2 Estimation of biological control prospects 

Weed Rating 



Bidens pilosa 



Cynodon dactylon 










Mimosa invisa 

Mimosa pigra 

Mimosa pudica 




Paspalum conjugatum 

PassiJlora foetida 






Any 

Family biological 

control 

successes? 

Asteraceae 

Amaranthaceae 

Asteraceae 

Asteraceae 

Commelinaceae 

Poaceae 

Poaceae 

Pontederiaceae 

Poaceae 

Euphorbiaceae 

Euphorbiaceae 

C yperaceae 

Marsileaceae 

Melastomataceae 

Asteraceae 

Mimosaceae 

Mimosaceae 

Mimosaceae 


Nephrolepidaceae 

Poaceae 

Poaceae 

Passifloraceae 

Poaceae 

Araceae 

Portulacaceae 

Poaceae 

Sphenocleaceae 

Attractiveness 

as a target in 

SE Asia 

unsuitable

4 Biological Control of Weeds: Southeast Asian Prospects 

3 Introduction 

Waterhouse (1 993a) published information, collated from agricultural and weed experts 

in the 10 countries of Southeast Asia, on the distribution and importance of their major 

weeds in agriculture. Ratings were supplied on the basis of a very simple system: 

+++ very widespread and very important 

++ not widespread but of great importance where it occurs 

+ important only locally 

present, but not an important pest 

The advantages and limitations of this system are discussed by Waterhouse (1993a). Of 

232 weeds nominated, 140 were rated as highly important, and a subset of 40 particularly 

SO. 

The focus of the present work is on the possibilities for classical biological control 

of those of this subset of 40 that evolved outside Southeast Asia. The assumption is that 

many of these have been introduced without some of the organisms that help to control 

them where they evolved. The chances are very remote indeed, for weeds that evolved in 

Southeast Asia, of introducing sufficiently host-specific organisms from outside the 

region. Nevertheless, it is possible that useful organisms present in, say, Thailand or 

Myanmar may not be present in all of the islands constituting the Philippines or 

Indonesia (or vice versa) and this possibility should be borne in mind. 

The origin of 12 of the subset of 40 major weeds is believed to be Southeast Asia, or 

close by, and these have been excluded from consideration at this stage. The remaining 

28 species, 27 of which are treated here, are either known to have evolved in the 

Americas or Africa or are postulated to have evolved in both Asia and Africa. This latter 

group is considered because the possibility exists that useful organisms at the African end 

of the range may not yet have extended their distribution into all of Southeast Asia. 

The 28th species, couch grass, Cynodon dactylon, has not been dealt with because, 

in many situations, such as lawns and some pastures, it is regarded as a highly desirable 

species. Biological control agents would not distinguish between these situations and the 

many others where it is a serious weed, so other control measures must be employed in 

the latter instances. 

Of course, it is not to be expected that all of any one country's top 20 or even top 10 

exotic weeds will necessarily be included in this regional priority list. Indeed, at least 

some of those omitted might well merit the production of additional dossiers if they are 

of such importance locally that resources for a program would be likely to achieve a very 

high priority for a particular country. ACIAR would be interested to hear of weeds that 

might be considered in this category. 

It is not so long ago that Wilson (1964) pointed out that no insects had yet been used 

for the biological control of aquatic weeds and that it was not clear "whether in the aquat- 

ic environment there exists a sufficient development of that monophagy in phytophagous 

insects that has been the main foundation for the biological control of weeds on land". He

Introduction 5 

referred to the opinion of Brues (1946) that aquatic insects show little host specificity, but 

warned that this view might be the result of lack of information and recommended an 

extension of research in this general field. In the intervening 30 years, research on four 

major water weeds of South American origin has yielded success and even spectacular 

success with the following: Salvinia molesta, Eichhornia crassipes, Alternanthera 

philoxeroides and Pistia stratiotes (Room 1993). 

It is very probable that a parallel can be drawn between the situation with water 

weeds in 1964 and the "conventional wisdom" of today that grassy weeds are unsuitable 

targets for classical biological control because of the danger to many major world crops 

that also belong to the family Poaceae e.g. rice, wheat, maize, sorghum, millet, sugar- 

cane. However, it would be very strange indeed if host specialisation occurred widely in 

insects attacking all other plant families, but not amongst those attacking the very large 

number of grasses. In view of the fact that 10 of the 18 world's worst weeds are grasses 

(Holm et al. 1977) and eight of the 28 major exotic weeds in Southeast Asia are also 

grasses (Waterhouse 1992, 1993a), it is evident that the time is long overdue for a 

detailed study of the natural enemies of these grasses in the regions where they evolved. 

This theme is mentioned further below, in particular in the discussion on Eleusine indica. 

For any biological control organisms to be approved for introduction into Southeast 

Asia against weedy grasses they would need to be sufficiently specific that they would 

not cause economic damage to the crop grasses listed in table 3.1. This list refers to 

Thailand, but is believed to be much the same as that for other Southeast Asian countries. 

It does not, however, include pasture species. A number of useful grasses are also har- 

vested from the wild and some may have to be considered also, although there are impor- 

tant weeds (e.g. Zmperata cylindrica) amongst them. There are, of course, many addition- 

al crop grasses of importance outside the region, but of little or no importance in most or 

all of Southeast Asia. They would certainly have to be taken into consideration in other 

regions of the world. 

The successful biological control of a weed presents a special problem, seldom 

shared by the control of an insect pest, namely that some other plant, perhaps even a 

weed that is more difficult to control by other means, will spread to occupy the space 

vacated. Reduction to the greatest possible extent of the density of a weed is desirable in 

situations such as pastures or national parks. In many other situations, however, all that 

may be required is a significant reduction in seeding (for annuals) or in competitiveness 

(for annuals and perennials) so that the weed no longer has an opportunity of becoming 

dominant and thus, when necessary, is more readily controlled by cultural or other mea- 

sures. Thus, even partial biological control (leading to the weed becoming less aggres- 

sive) provides desirable plant species with the opportunity to compete more successfully 

for sunlight and nutrients and may be of significant value. 

Another problem is that many weeds display a good deal of variability throughout 

their distribution, resulting in part from polyploidy, hybridisation with closely related 

species and other genetic modifications. The taxa thus produced may not be equally sus- 

ceptible to natural enemies, so it is desirable, where possible, to match-tb-wlth ixa 

encountered in the surveys in the area of origin of the weed. It may also be necessary to


seek expert taxonomic advice at an early stage, perhaps involving electrophoretic, DNA 

and other studies, particularly when commencing a project on a weed that has not yet 

been the target of a biological control investigation. 

The summary accounts presented are designed to enable a rapid review to be made 

of (i) the main characteristics of the major weeds of agriculture that are believed to be 

exotic to part or all of Southeast Asia, (ii) what is known of their natural enemies and 

(iii) prospects for reducing their weediness by classical biological control. 

The material on weed characteristics draws heavily on the publications by Barnes 

and Chan (1990), Holm et al. (1977), Noda et al. (1985) and Soerjani et al. (1987). 

Additional information is available from these sources, including detailed botanical 

descriptions, vernacular names, biology, agricultural importance and herbicidal control. 

I am particularly grateful to the University of Hawaii Press for permission to draw 

on 21 of the illustrations in its publication 'The World's Worst Weeds' by Holm et al. 

(1977) to Ancom Berhad, Malaysia (Barnes and Chan 1990) and the Director of 

BIOTROP Indonesia (Soerjani et al. 1987) to draw on 2 and 3 illustrations respectively 

from their publications and to the Division of Entomology CSIRO for permission to use 

illustration 4.16. The figures have been slightly amended by the omission of inserts that 

are mainly of taxonomic interest. Acknowledgement appears on each of the illustrations 

used. 

In most instances four databases were searched for relevant information: 

AGRICOLA (Bibliography of Agriculture) 1 970+ 

BIOSIS (Biological Abstracts) 1989+ 

CAB (Commonwealth Agricultural Bureaux) 1984+ 

DIALOG (Biological Abstracts) 1959+ 

1 many cases abstracting journals and other sources published prior to the above 

cominencement dates were also searched. Useful information was also obtained by 

serendipity from these and other references and from unpublished records. Nevertheless, 

in many cases the search cannot be described as exhaustive. Even more relevant, howev- 

er, than attempting an exhaustive search would be a fresh, detailed field survey targeted 

on the known (or presumed) area of origin of the weed. In any event, in most instances a 

preliminary investigation would be highly desirable in the area of origin of a weed before 

deciding whether or not to embark upon a major project. Several such surveys might well 

be carried out simultaneously where more than one weed occurs in the same general 

region. Indeed, it is strongly recommended that a pre-project activity be funded to carry 

out such surveys, with special reference to selected weeds of major importance in 

Southeast Asia. 

Surveys of this nature are particularly important, since the amount of useful, pub- 

lished information on arthropods or other organisms attacking the target weeds is, in gen- 

eral, inadequate to serve as a basis for a sound decision. Although acceptable host speci- 

ficity is required for classical biological control, it is possible that some of the less specif- 

ic fungi listed might be developed for use as bioherbicides. 

In addition to surveys in the region of origin of the weed(s) it will also be necessary 

to survey the weed(s) in the country or countries where biological control is to be

Introduction 7 

attempted. This is to indicate whether any of the organisms that might be considered for 

introduction are already present. 

The species treated are drawn from tables 10 and 11 of 'The Major Arthropod Pests 

and Weeds of Agriculture in Southeast Asia: Distribution, Importance and Origin' 

(Waterhouse 1993a). It is quite possible that additional weeds rating highly in these 

tables will prove to be exotic to Southeast Asia (or significant parts of it) and, alternative- 

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

in the region. 

The natural enemies most commonly involved in classical biological control of 

weeds have been arthropods, although there is a growing interest in, and a few striking 

successes with, fungi. Because there is a considerable lack of uniformity in the names of 

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

names. These have been used in the text, replacing those used by the authors quoted. On 

the other hand, with few exceptions the names used for fungi, bacteria, nematodes and 

viruses are those of the authors quoted, although it is probable that some names have 

been changed since they were used. Where the name of a weed or an insect given 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 Dr B. 

Napompeth (Thailand), Dr R. Muniappan (Guam), C.J. Davis (Hawaii) and, in Australia, 

Dr I.W. Forno, Dr K.L.S. Harley, M.H. Julien, Dr K.R. Norris, J. Prance, Dr D.P.A. 

Sands, Dr A.J. Wapshere and A.D. Wright of CSIRO and Dr R.E. McFadyen 

(Queensland Department of Lands). Many others who have contributed unpublished 

information are acknowledged at appropriate places in the text. 

Valuable advice on taxonomic problems has been received from a number of col- 

leagues in the Division of Entomology, CSIRO, Canberra, including Dr M. Carver 

(Hemiptera), Dr P. Cranston (Diptera), E.D. Edwards (Lepidoptera), Dr I.D. Naumann, 

Dr K.H.L. Key (Orthoptera), T. Weir (Coleoptera) and Dr E.C. Zimmerman 

(Curculionidae). 

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

Coordinator, Crop Scienqs, ACIAR, Canberra. 

It is a pleasure to acknowledge the expert assistance of Mrs A. Johnstone 

(Ms A. Ankers) in converting my manuscripts into presentable form; and also of Mrs S. 

Smith and C. Hunt for assistance with the illustrations. 

It would not have been possible to continue with these biological control activities 

in deep retirement without the support, forbearance and encouragement of my wife, to 

whom particular thanks are due.

8 

Biological Control of Weeds: Southeast Asian Prospects 

Table 3.1 Grasses (other than pasture species) that are important in Thailand. 

A. Crop Grasses Importance 

Bambusa spp. 

Coix lacryma-jobi 

Cymbopogon spp. 

Hordeum spp. 

Oryza sativa 

Saccharum oficinarum 

Setaria italica 

Sorghum bicolor 

Triticum spp. 

Zea mays 

Zizania latifolia 

B. Grasses harvested from the wild 

Arundo donax 

Dendrocalamus spp. 

Gigantochloa spp. 

Imperata cylindrica 

Melocanna baccifera 

Phragmites spp. 

Phyllostachys spp. 

Schizostachyum dumetorum 

bamboo, construction, furniture, 

paper 

job's tears, cereal 

lemongrasses, flavourings 

barleys 

rice 

sugar cane 

foxtail millet 

sorghum 

wheats 

maize 

vegetable 

giant reed, cane 

weaving, vegetables 

construction, furniture 

paper, roof thatch 

paper, furniture, food 

reeds, thatch, mats 

furniture, vegetable 

rope

Target weeds 



Bidens pilosa 












Mimosa invisa 

Mimosa pigra 

Mimosa pudica 



Panicurn repens 

Paspalum conjugatum 

Passiflora foetida 

Penniseturn polystachion 




Sphenoclea zeylanica



(after Holm et a/. 1977)

Map 4.1 Ageratum conyzoides 

4.1 Ageratum conyzoides 11 


As a member of the Asteraceae, it would be expected that Ageratum conyzoides would 

have many natural enemies attacking it in its area of origin in Tropical America. 

However, no study has been made and virtually nothing is known of the situation there. 

Elsewhere it is attacked by a range of insects, nematodes, fungi and viruses, but almost 

all have a very wide host range and are not suitable as biological control agents. 

Surveys in Tropical America would be necessary to provide data on which 

prospects for its biological control could be evaluated.


4.1 Ageratum conyzoides L. 

Asteraceae 

goatweed, ageratum; bandotan (Indonesia), rumput tahi ayam (Malaysia), bulak 

manok, kolokong kabayo (Philippines), ya tabsua, ya sap raeng (Thailand), 

co c~lt heo, bo xit (Vietnam) 

Rating 

+++ Myan, Thai 

17 ++ Msia, Sing, Phil 

+ Laos, Camb, Viet, Brun, Indo 

Origin 

Tropical America. 

Distribution 

Pantropical; also in the subtropics and extending into temperate areas from latitude 30°N 

to 30's. Widespread in SE Asia. Present in Java prior to 1860. 

Characteristics 

Ageratum conyzoides is a self pollinated, C3, annual herb. It is erect, often branched, 

sometimes decumbent and ranges up to 1.2 m at flowering. Its flowers are light blue, 

white or violet and its leaves and stems are hairy. 

Importance 

A. conyzoides occurs in both light and heavy soils in moister areas in agricultural land, 

waste land, roadsides, plantations, pastures and upland rice fields. It may produce 

40 000 or more seeds per plant and these are mainly spread by wind and water. They 

germinate readily and the life cycle can be completed in less than 2 months. A. cony- 

zoides is one of about 300 species in the genus, all of which originated in the Americas. 

Goatweed is important in 46 countries in 36 crops and is troublesome in plantations 

after grasses have been suppressed (Holm et al. 1977). It is a rapidly colonising, vigor- 

ously growing weed in a wide variety of arable crops in which thick carpets of A. cony- 

zoides compete strongly for nutrients and moisture. When a stand is destroyed another 

rapidly takes its place. It is suspected of poisoning cattle, but this is not confirmed from 

Australia. It was rated 19th of the World's Worst Weeds by Holm et al. (1977), as equal 

15th in Southeast Asia (Waterhouse 1993a) and 15th in the Oceanic Pacific 

(waterhouse unpub.). 

Its crushed leaves smell strongly of coumarin and are used as a styptic for wounds, 

also for sores, skin diseases, eye inflammation and lung problems (Gonzalez et al. 1991). 

It is sometimes used as cut flowers in the home.

Natural enemies 


Although A. conyzoides is listed by Holm et al. (1977) as a weed in some crops in 

Central and South America, it is significant that nowhere in that region (unlike the rest of 

the tropical world) is it regarded as a serious or a principal weed. From this it might be 

inferred that natural enemies might be controlling its abundance. However, so little infor- 

mation on natural enemies (Tables 4.1.1 to 4.1.3) was obtained from the databases 

searched that it is not possible to substantiate this claim. Almost all of the records are 

from outside its area of origin and one (the agromyzid fly, Melanagromyza metallica) is 

known to have a narrow host range. However, M. metallica is already widespread. In 

addition to India, it is known also from many places including Taiwan, Philippines, 

Vietnam, Thailand, Indonesia, Melanesia, Papua New Guinea, Solomon Is, Micronesia, 

Australia and Africa. It lays its eggs on the apical part of the stem. The larva bores into 

the pith region, gradually extending towards the root and the final instar larva cuts an exit 

hole at the base of the stem. Mines may extend into the roots and pupae are often present 

at about ground level in the mines (Singh and Beri 1973). 

If A. conyzoides is considered to be an important target it will be necessary to sur- 

vey for organisms attacking it in Central America and northern South America. 

Table 4.1.1 Natural enemies of Aaeratum convzoides: insects and mites. 

Species Location Other hosts References 

INSECTS 

Orthoptera 

ACRlDlDAE 

Zonocerus Nigeria 

variegatus 

Hemiptera 

APHlDlDAE 

Aphis craccivora 

Aphis gossypii 

Aphis spiraecola Java 

(= A. nigricauda) 

Aulacorthum 

solani 

Brachycaudus 

helichrysi 

Capitophorus 

hippophaes 

Hyperomyzus 

carduellinus 

Myzus ormtus 

Myzus persicae 

Neomasonaphis 

(- Masonaphis) 

amphalidis 

Uroleucon 

(= Macrosiphum) 

solidaginis 

Vesiculaphis pieridis India 

Chromolaena odorata, 

Lantana camara 

many 

many 

many 

many 

many 

Eupatorium, Mirabilis, 

Polygonum 

many 

many 

many 

Lyonia ovalifolia, 

Pieris ovalifolia 

Raychaudhuri 1983 


Patch 1939, 




Ghosh et al. 1971 

Patch 1939 




Patch 1939 

Patch 1939 

(continued on next page)


Table 4.1.1 (continued) 


ALEYRODIDAE 

Bemisia tabaci 

DlASPlDlDAE 

Mycetaspis personata 

LYGAEIDAE 

Nysius inconspicuus 

Thysanoptera 

PHLAEOTHRIPIDAE 

Haplothrips gowdei 

THRlPlDAE 

Calipthrips ipomoeae 

Microcephalothrips 

abdominalis 

Thrips tabaci 

Diptera 

AGROMYZIDAE 

Calycomyza sp. 

Melanagromyza 

metallica 

CHLOROPIDAE 

Olcella pleuralis 

TEPHRITIDAE 

Xanthaciura imecta 

Lepidoptera 

ARCTll DAE 

Pareuchaetes 

pseudoinsulata 

(= Ammalo insulata) 

GELECHllDAE 

Dichomeris sp. 

NOCTUIDAE 

Pseudoplusia includem 

(=Plusia 00) 

Spodoptera frugiperda 

India, a very wide range Ang et al. 1977, 

Malaysia, Sastry 1984, 

Turkey Shreni et al. 1979 

Brazil polyphagous dlAraujo e Silva et al. 1968a 

India sesame and many Thangavelu 1978 

others 

Hawaii vector of pineapple Sakimura 1937 

yellow spot virus 

Brazil polyphagous dlAraujo e Silva et al. 1968a 

India polyphagous Gopinathan et al. 198 1 

Hawaii vector of pineapple Sakimura 1937 

yellow spot virus 

USA Spencer & Steyskal 1986 

India, etc no other host Singh & Beri 1973 

mentioned 

Trinidad C. odorata, C. ivaefolia, McFadyen 1988a 

C. iresinoides, 

Fleischmannia 

microstemon, 

Wedelia 

caracasana, 

Wulfla baccata 

Florida, C. odorata, 

Trinidad F. microsternon 

W. caracasana 

McFadyen 1988a, 

Needham 1946 

Nigeria, C. odorata Bennett & Cruttwell 1973, 

Trinidad Olaoye 1974 

Trinidad C. odorata Bennett & Cruttwell 1973 

Brazil polyphagous d'Araujo e Silva et al. 1968a 

Brazil polyphagous d'Araujo e Silva et al. 1968a 


MITES 



PYRALIDAE 

Pionea upalusalis Trinidad, C. odorata, C. ivaefolia, McFadyen 1988a 

Puerto Rico, Austroeupatorium 

Venezuela inulaefolium, 

Fleischmannia 

microstemon 

Brevipalpus obovatus India cotton, Solanum nigrum, Sadana et al. 1983 

Sonchus asper, 

Phaseolus vulgaris, 

Euphorbia hirta, 

Xanthium sp., 

Cichorium intybus 

Tetranychus urticae China a very wide range Dong et al. 1986 

Table 4.1.2 Natural enemies of Ageratum conzoides: nematodes. 


Aphelenchoidesfiagariae 

Helicotylenchus multicinctus 

Meloidogyne sp. 

Meloidogyne arenaria 


thamesis 

Meloidogyne incognita 

Meloidogyne javanica 

Pratylenchus pratensis 

Rotylenchulus reniformis 

Hawaii strawberry, Vanda 

orchids, Impatiens, 


Brazil banana, Portulaca 

oleracea and several 

weeds 

Cuba Eleusine indica, 

Croton lobatus, 


Philippines 

Philippines 

Philippines many vegetables 

and weeds 

Philippines, many vegetables 

Nigeria and weeds 

Hawaii 

Hawaii, India many weeds 

Sher 1954 

Zem & Lordello 1983 

Acosta et al. 1986 

Holm et al. 1977 

Valdez 1968 

Valdez 1968 

Mamaril& Alberto 1989 

Mamaril& Alberto 1989, 

-Salawu et al. 1991 

Valdez 1968 


Linford & Yap 1940, 

La1 et al. 1978


Table 4.1.3 Natural enemies of Ageratum conyzoides: fungi, bacteria and viruses. 


FUNGI 

Cercospora agerati 

Colletotrichium sp. 

Cylindrocladium 

quinqueseptalum 

Mycovellosiella perfoliata 

Puccinia conoclinii 

Sclerotium rolfsii 

BACTERIA 

Pseudomonas 

solanacearum 

India 

India 

India 

India 

many commercial 

hosts 

many 

India potato, Ranunculus 

sceleratus 

Stevens 1925 

Kulkami & Sharma 1976 

Sulochana et al. 1982 

Srivastava 198 1 

Stevens 1925 

Desai et al. 1980 

Sathiarajan & 

Sasikumar 1977, 

Sunaina et al. 1989 

VIRUSES 

Ageratum vein yellowing India, (transmitted by 

Ang et al. 1977, 

Malaysia Bemisia tabaci) 

Shreni et al. 1979 

anemone mosaic 


Bidens mottle several, including 

Zinnia, petunia 

& Verbena 

Logan & Zettler 1984 

hibiscus yellow vein India (transmitted by 

Jeyarajan et al. 1988 

mosaic 

B. tabaci) 

pineapple yellow spot Hawaii 

Sakimura 1937 

potato virus Y 

India 

potato 

Joshi & Prakash 1977 

tapioca mosaic 

India 

(transmitted by 

B. tabaci) 


tobacco leaf curl India tomato 

Holm et al. 1977, 

Reddy et al. 1981 

tomato leaf curl Turkey, many weeds 

Sastry 1984, 

India 

(transmitted by 

B. tabaci) 


urd bean yellow mosaic India (transmitted by 

B. tabaci) 


Zinnia yellow net India (transmitted by 

B. tabaci) 

Srivastava et al. 1977




Map 4.2 Amaranthus spinosus 

4.2 Amaranthus spinosus 19 

Amaran thus spinosus 

Mass rearing and release, as required, of the weevil Hypolixus trunculatus is reported to 

provide good control of Amaranthus spinosus in Thailand but, of course, this is augmen- 

tative rather than classical biological control. 

Three other insects (a weevil, a leaf mining fly and a caterpillar) are known which may 

prove to be adequately specific for classical biological control. 

However, almost nothing is known about the natural enemies of A. spinosus in tropical 

America where it evolved and it would thus be necessary to cany out a survey there in 

order to evaluate what potential biological control agents are available.


4.2 Amaranthus spinosus L. 

Amaranthaceae 

spiny amaranth, spiny pigweed, needle burr; hin nu nive tsu bauk (Myanmar), 

phak khom nam (Thailand), phti banla (Cambodia), bayam duri (Malaysia and 

Indonesia), orai (Philippines), den gai (Vietnam) 

Rating 

+++ Myan, Thai, Phil 

17 ++ Msia, Sing 

+ Laos, Camb, Viet, Indo 

Origin 


Distribution 

A. spinosus is mainly tropical and subtropical in distribution, but also extends into the 

temperate zone from latitude 30°N to 30"s. 


A. spinosus is an erect, much branched, annual, growing to 1.2 m. Its stems are angled in 

cross section, fleshy, often reddish and bear many spines. Its leaves are alternate, with a 

pair of straight spines up to 1 cm long at the base. The inflorescence is long, slender and 

terminal or arises from leaf axils. The flowers are small, greenish and unisex. It is propa- 

gated by reddish brown seeds. 

Importance 

Spiny amaranth prospers in warm sunny situations, but not where it is cool or shady. It is 

not reported as a problem in the Mediterranean or Middle East. It is a weed in 44 coun- 

tries in 28 crops, mainly in the Caribbean, in the west and south of Africa, in India and in 

Southeast Asia. Up to 235 000 seeds per plant have been recorded. Seeds are spread by 

wind and water. Some germinate soon, others over several months and still others remain 

viable in the soil for many years. A. spinosus is abundant in cultivated and abandoned 

fields, along roadsides and in waste places. It is a weed of varying degrees of aggressive- 

ness in many crops, including upland rice, cotton, cowpeas, groundnuts, maize, mangos, 

millet, pineapples, sugarcane and vegetables. The rigid needle-like spines break off in the 

hands of workers in sugarcane, cotton and other crops. 

A. spinosus may contain high nitrate levels and has been implicated in livestock poi- 

soning. It is avoided by most animals because of its spines. Leaves are sometimes used 

by humans as a green vegetable. Other Amaranthus species are valuable as a grain crop 

in some South American countries and the family Amaranthaceae contains a number of 

widely grown ornamental garden species (Purseglove 1968).



A. spinosus is attacked by a number of natural enemies (Tables 4.2.1 and 4.2.2), but most 

of the reports come from outside its native range and are of non-specific organisms. The 

agromyzid fly Haplopeodes minutus, known in USA from species of Amaranthus and 

Chenopodium (Spencer and Steyskal 1986) and both the beetle Cassida nigriventris and 

the moth Coleophora versurella, known in Pakistan from these same plant genera (Khan 

et al. 1978), may prove to be sufficiently specific to be candidate biological control 

agents. 

The weevil Hypolixus trunculatus, whose larvae tunnel in the stems and form galls, 

is known from Pakistan, India and Thailand and attacks Amaranthus spinosus, A. viridis 

and Digera arvensis. Although it has a relatively long life'cycle and low reproductive 

capacity, mass rearing and augmentative releases have resulted in a satisfactory level of 

control and replaced the use of herbicides in Thailand (Julien 1992, Napompeth 1982, 

1989, 1992a). Females deposit eggs singly in cavities scooped out of the shoots. Larvae 

tunnel down inside the stem to its base, where a gall develops. Breeding continues 

throughout the year but is at its height in late summer. At this time the life cycle is 44 to 

50 days. Pupation occurs within the gall. Larvae and pupae are parasitised by larvae of 

the pteromalid wasp Oxysychus sp. (Aganval 1985). 

Evans (1987) records five fungi from A. spinosus but, except for one which is 

unsuitable because it has a wide host range, too little is known about their host specificity 

to assess the prospects for their use in classical biological control. 

Comment 

Almost nothing is known about the natural enemies of A. spinosus in tropical America 

where it evolved. A survey in this region would be necessary to document the organisms 

attacking it. There are good general grounds for believing that there are some natural 

enemies that are specific to the family Amaranthaceae. In most countries, members of 

this family have little value as crop plants, so the chances are that some safe natural ene- 

mies will be found that are of value as classical biological control agents. 

Table 4.2.1 Natural enemies of Amaranthus spinosus: insects and mites. 


INSECTS 

Hemiptera 

APHlDlDAE 

Myzus persicae Malawi, highly polyphagous Chapola 1980, Napompeth 

Thailand 1982 

COREIDAE 

CIetus fuscescens Nigeria Amaranthus dubius, Ukwela & Ewete 1989 

A. cruenrus, 

A. hypochondriachus 

LYGAEIDAE 

Germalus unipunctarus Vanuatu Cock 19841, 

Nysius sp. Vanuatu Cock 1984b 



Table 4.2.1 continued 


MlRlDAE 

Horcias nobilellus Brazil ' polyphagous d'Araujo e Silva et al. 1968a 

PI ESMATIDAE 

Piesma cinereum Brazil polyphagous d'Araujo e Silva et al. 1968a 

Thysanoptera 


Haplothrips India Amaranthus viridis, Dhiman 1986 

longisetosus A. oleosa, Chenopodium 

anthelminthicum 

Coleoptera 

CHRYSOMELIDAE 

Cassida exilis 

Cassida nigriventris 

CURCULIONIDAE 

Ceutorhynchus 

asperulus 

Hypolixus 

trunculatus 

MELYRIDAE 

Astylus lineatus 

Pakistan Amaranthus viridis, 

Chenopodium album 


Chenopodium album, 

Spinucia oleracea 

India red gram, Amaranthus 

viridis, A. tricolor, 

Basella alba 

Pakistan, Amaranthus viridis, 

India, Chromolaena odorata 

Thailand Digera arvensis 

Brazil citrus 

Baloch et al. 1976 


Khan et al. 1978 

Puttaswamy & 

Channabasavannal98 1, 

Puttaswamy et al. 198 1 

Aganval 1985, 

Baloch et al. 1976, 1977, 

Ghani 1965, Julien 1992 

Napompeth 1982, 1990b, 

1992a 

d'Araujo e Silva et al. 1968a 

Diptera 

AGROMYZIDAE 

Haplopeodes USA Amaranthus, Spencer & Steyskal 1986 

minutus Chenopodium 

Lepidoptera 

COLEOPHORIDAE 

Coleophora 

versurella 

CURCULIONIDAE 

Hypolixus ritsemae 

LYCAENIDAE 

Zizeeria knysna 

Zizeeria krupta 

NOCTUIDAE 

Neogalea 

(= Spodoptera) sunia 

Spodoptera eridania 

Spodoptera exigua 

. Spodoptera litura 

1 

Pakistan 

Vanuatu 

Pakistan 

Pakistan 

Nicaragua 

Nicaragua 

Nicaragua 

Philippines 

Chenopodium botrys Khan et al. 1978 

Cock 19841, 


Baloch et al. 1977, Ghani 

1965 

polyphagous Savoie 1988 



highly polyphagous Moody et al. 1987 




PYRALI DAE 

Loxostege sp. Argentina seed heads of 

Amaranthus sp. 

(the genus Loxostege 

Spoladea (=Hymenia) 

recurvalis 

SCYTHRIDIDAE 

Eretmocera 

impactella 

TORTRICIDAE 

Archips sp. 

YPONOMEUTIDAE 

Plutella xylostella 

MITE 

TETRANYCHIDAE 

Tetranychus 

novocaledonicus 

India, 

Pakistan 

Vanuatu 

contains pests) 

polyphagous 


Chenopodium album 

Pakistan 

Pakistan 

India Amaranthus tricolor, 

4. viridis 

C.J. Deloach 

pers. comm. 1980 


Chaudhury & Kapil 1977, 

Lock 1984b. Ghani 1965 


Ghani 1965 

Ghani 1965 

Puttaswav.y dr 

Channabasavanna 1981 

Table 4.2.2 Natural enemies of Amaranthus spinosus: nematodes, fungi, viruses. 


NEMATODES 

Cactodera amaranthi Cuba spinach, other species Stoyanov 1972 


Pratylenchus zeae 

Pseudocephalobus indicus 

Philippines 

of Amaranthus 

rice, many weeds 

India only recorded on 

Valdez 1968 

Fortuner 1976 

Joshi 1972 

Rotylenchulus reniformis India, USA 

A. spinosus 

many weed hosts Inserra et al. 1989, 

La1 et al. 1978 

FUNGI 

Albugo bliti Dominica, many Amaranthaceae 

Jamaica, India, 

Pakistan, Sudan 

Alternaria compacta 

Aposphaeria amaranthi 

Bipolaris indica (as 

Drechslera indica) 

India 

USA potential bioherbicide 

for A. albus; effect on 

A. spinosus not known 

many, including 

Helianthus, Pennisetum, 

Portulaca 

Baloch et al. 1977, 

Evans 1987 

Kar & Ashok-Das 1988 

Mintz & Weidemann 1992 

Evans 1987, 

Kenfield et al. 

1989 



Table 4.2.2 continued 


Cercospora brachiata 

(= C. amaranthi) 

Fusarium oxysporum f.sp. 

elaeidis 

Phoma tropica 

Puccinia sp. 

VIRUSES 

cucumber mosaic 

Digera mosaic 

groundnut rosette 

tobacco bunchy top 

tobacco mosaic 

India, Nigeria, many Amaranthaceae 

Uganda, Trinidad, 

USA, Japan, 

China, USSR 

Nigeria oil palm, Chromolaem 

odorata, Impera ta 

cylindrica, Mariscus 

alternifolius 

India 

Hong Kong 

India cucumber, Solanum, 

nigrum, Tagetes 

erecta, etc 

India several weeds 

Malawi (Myzus persicae 

Philippines is a vector) 

Evans 1987 

Oritsejafor 1986 

Evans 1987 

Evans 1987 

Suteri et al. 1980 

Singh et al. 1975 

Adams 1967 

Chapola 1980 

Eugenio & del Rosario 1962


Bidens pilosa 


Map 4.3 Bidens pilosa 

4.3 Bidens pilosa 

Bidens pilosa is native to tropical America. Preliminary studies, based mainly on 

Trinidad, indicate that it is attacked by a number of natural enemies, mainly insects, and 

that several of these may be sufficiently host specific to be considered as biological con- 

trol agents. Further host specificity studies are required and additional, wider-ranging 

searches, particularly in South America.


4.3 Bidens pilosa L. 

Asteraceae 

cobbler's pegs, Spanish needle; djaringan ketul (Indonesia), pisau pisau 

(Philippines) yah koen jam khao (Thailand) 

Rating 

++ Thai, Indo, Phil 

10 + Myan, Laos, Camb, Viet 

Msia 

Origin 

Tropical America 

Distribution 

Pantropical. Known from Java before 1835, but apparently not present in Kalimantan or 

the Moluccas (Soerjani et al. 1987). 


Bidens pilosa is an erect, slender, branching, annual herb growing up to 1.5 m. Its stems 

are four-angled in cross section and its leaves opposite and sparsely hairy. The abundant 

yellow flowers are borne in heads on long stalks and produce black, barbed seeds charac- 

teristically radiating in all directions from a common base. The recurved, 2-toothed barbs 

enable the seeds to stick readily to hair and clothing and they are also distributed by wind 

and water. Cobbler's pegs prefers moister soils and flowers all year round. 

Importance 

A very common weed of 31 crops in more than 40 countries, B. pilosa occurs in gardens, 

cultivated land, open waste places and along roadsides. It is an important weed of pas- 

tures, maize, sorghum, vegetables, cotton, tea, coffee, cassava, coconut, oil palm, citrus, 

papaya, rice, rubber and tobacco. Single plants produce up to 6000 seeds, many of which 

germinate readily, permitting three or four generations a year in some regions. 

Some seeds remain viable in the soil for at least 5 years. When herbicides have 

eradicated perennial grasses this weed often becomes dominant. 

In South Africa the early spring growth is sometimes eaten by humans, but has low 

nutritive value. It has a pungent essential oil that may taint milk. 


These 'are also dealt with in 'Biological Control: Pacific Prospects' (Waterhouse and 

Norris 1987) which did not assess B. pilosa a particularly promising target for biological 

control. However, more information has since become available (Table 4.3.1 and 4.3.2),

4.3 Bidens pilosa 29 

particularly concerning leaf miners and seed head feeders of the fly family Agromyzidae. 

This suggests that there may be good prospects for some of these natural enemies. 

Few details are available of the natural enemies of B. pilosa in Brazil. The pupal 

stage of the chrysomelid beetle Phaedon pertinax (= P. consimilis) lasts 6 to 8 days and 

the pentatomid bug Stiretrus erythrocephalus passes through 4 instars in 30 days 

(Ribeiro 1953). Thrips killed 22.25% of B. pilosa plants (particularly seedlings) and 

Diptera infested 97.8% of flower heads. Parasitisation of these Diptera by wasps and 

flies, varied from 40.96% to 58.91 % according to the size of the population (Esposito et 

al. 1985). 

About half of the 2500 species of the family Agromyzidae have known hosts and 

almost all of this group are restricted in their feeding to a single family or genus. Only 16 

of the species (0.6% of the total) are truly polyphagous, feeding on a number of unrelated 

families (Spencer 1990). Agromyzid flies are, therefore, worth serious consideration as 

classical biological control agents. In this context, plants of the genus Bidens appear to be 

particularly attractive to agromyzid flies for they support 19 species (Table 4.3.3). 

In the tribe Coreopsideae (of the family Asteraceae) only two (Bidens and 

Coreopsis) of its 26 genera support Agromyzidae (Table 4.3.4). Coreopsis is native in 

North America, but no agromyzids are known on it there, although three polyphagous 

species are known to attack it in Europe, India or Australia (Spencer 1990). 

Eleven of the above 19 species are known from Bidens pilosa (Table 4.3.1). Of 

these, three are restricted to the genus Bidens (perhaps even to B. pilosa), two are 

polyphagous, and the remaining six have one or more additional hosts in other genera of 

the Asteraceae. Ten of the eleven species are restricted to the Americas and further host 

specificity tests may well indicate that many are valuable biological control agents. Four 

of the ten form blotch mines (Arnauromyza maculosa, Calycomyza allecta, C. platyptera 

and Liriomyza archboldi), one makes long, linear irregular mines (Liriomyza venegasiae), 

and three feed in the seed heads (Liriomyza insignis, Melanagromyza bidentis and M. 

floris) (Spencer 1990, Spencer and Steyskal 1986). 

The flower heads of B. pilosa are also attacked by three species of Tephritidae in 

Central America and by one of these in India. Adult weevils of the genera Baris, 

Centrinaspis and Promecops feed in the flowers of B. pilosa and other Asteraceae, but 

are thought not to breed there. Several other insects (at least three other beetles and a 

pierid butterfly) have also been recorded from B. pilosa and sometimes from other 

Asteraceae as well. 

Table 4.3.4 shows the position of the genus Bidens as a member of the tribe 

Coreopsidae, within the family Asteraceae. There may well be natural enemies that 

attack it, but not any species of agricultural or special environmental significance. 

Attempts at biological control 

There have been none.


Table 4.3.1 Natural enemies of Bidens pilosa: insects. 


Hemiptera 

ALEYRODI DAE 

Dialeurodes vulgaris India coffee, Erythrina 

lithosperma 

Venkataramaiah 1974 

APHlDlDAE 

Aphis coreopsidis Brazil soybean Almeida 1979, d'Araujo 

e Silva et al. 1968a 


Christie et al. 1974, 


Aphis illinoisensis Brazil 

Uroleucon 

(= Dacrynotus) sp. 

MlRlDAE 

Brazil, USA tobacco, lettuce 

Garcanus gracilentus Brazil sweet potato, 

polyphagous 


Horcias nobilellus 

PENTATOMIDAE 

Brazil polyphagous 



Stiretrus erythrocephalus Brazil 

Ribeiro 1953 

Thyanta perditor Brazil soybean 

Grazia et al. 1982 

Coleoptera 

APlONlDAE 

Apion luteirostre 

CHRYSOMELIDAE 

Chalcophana viridipennis 

Chlamisus insularis 

Phaedon pertinax 

(= P. consimilis) 

Physimerus pygmaeus 

CURCULIONIDAE 

Baris sp. 

South America Mikania micrantha Cock 1980 

Brazil 

Trinidad Chromolaena odorata, 

C. ivaefolia 

Brazil, Mikania micrantha 

(not in Trinidad) 

South America Mikania micrantha 


McFadyen 1988a 

Cock 1980, d'Araujo e Silva 

et a]. 1968a, Ribeiro 1953 

Cock 1980 

Trinidad (feed in B. pilosa 

flowers) 

Cruttwell 1971a 

Centrinaspis sp. Trinidad (feed in B. pilosa 

flowers) 

Cruttwell 197 1 a 

Promecops sp. Trinidad (feed in B. pilosa 

flowers) 

Cruttwell 197 la 

Rhodobaenus 

Trinidad adults feed on stems, McFadyen 1988a 

cariniventris 

and petioles of B. pilosa, 


C. ivaefolia, 

Austroeupatorium 

inulaefolium 

Rhodobaenus, 

Trinidad feed in B. pilosa stems: McFadyen 1988a 

tredecimpunctatus 

and in several other 

Asteraceae 

Diptera 

AGROMYZIDAE 

Amauromyza maculosa Trinidad (also polyphagous, but favours 

N&S America, Asteraceae 

Hawaii) 

Cruttwell 197 1 a, 

Spencer 1990, Spencer & 

Steyskal 1986 




Calycomyza allecta Trinidad (also 

Brazil, 

Guadeloupe, 

Venezuela) 

Calycomyza USA (Florida, 

platyptera California) 

Liriomyza archboldi Florida 

(Bahamas, 

Costa Rica) 

Liriomyza insignis Costa Rica 

Liriomyza trifolii cosmopolitan 

Liriomyza venegasiae Southern 

California 

Liriomyza sp. Argentina 

Melanagromyza bidentis Florida, 

Caribbean 

Melanagromyzafloris Costa Rica, 

Mexico, 

Trinidad 

(also Florida, 

Neotropics) 

Melanagromyza splendid USA, Hawaii 

Phytomyza atricornis Australia 

CEClDOMYllDAE 

Asphondylia bidens Florida 

DROSOPHILIDAE 

Cladochaeta nebulosa Florida 

TEPHRlTlDAE 

Dioxyna sororcula Florida, 

(= D. picciola) Trinidad, 

widespread 

Xanthaciura insecta Florida, 

Trinidad 

Lepidoptera 

ARCTllDAE 

Hypercompe Brazil 

(=Ecpantheria) 

hambletoni 

NOCTUIDAE 

Cropia (=Dyops) minthe Brazil 

Mocis latipes Brazil 

Thysanoplusia Kenya 

(- Diachrysia) orichalcea 

PlERlDAE 

Perrhybris phaloe Trinidad 

(= Ascia buniae phaloe) 

Helianthus, Rudbeckia 

and garden Asteraceae 

Asteraceae, including 

Aster, Helianthus, 

Zinnia 

restricted to Bidens 


polyphagous, including 

Chrysanthemum 

Venegasia carpesioides 


Verbesina sp. 

Calendula sp 

Asteraceae including 

Helianthus, Lactuca 

polyphagous, including 

Cineraria 

attacks several 

Asteraceae in India 

Ageratum conyzoides, 


Fleischmannia 

caracasana 

Panicum maximum, 

Paspalum notarum, 

Hyparrhenia rufa 

coffee and other crops 

Cruttwell 1971b, Frick 

19.56, Spencer 1990, 

Spencer & Steyskal 1986 

Spencer 1990 


Spencer 1990, 


Spencer 1990 

Spencer 1990, 


Spencer 1990 

Spencer 1990 

Cruttwell 197 la, Spencer 

1990, Spencer & 

Steyskal 1986 

Spencer 1990, 


Kleinschmidt 1970 

Steyskal 1972 

Steyskal 1972 

Cruttwell 1971 a, 1972a,b, 

Steyskal 1972 

McFadyen 1988a, 

Steyskal 1972 



Lourencao et al. 1982 

Bardner & Mathenge 1974 

Cruttwell 197 1 a

32 


Table 4.3.2 Natural enemies of Bidens pilosa: nematodes, fungi, mycoplasmas, 

viruses. 

Species Location 

NEMATODES 

Meloidogyne sp. Hawaii 

Meloidogyne hapla India 

Pratylenchus minutus Hawaii 

Rotylenchulus reniformis USA 

FUNGI 

Cercospora bidenfis Mauritius 

Cercospora rnegalopotamica Hawaii 

Entyloma guaraniticurn Mauritius 

Uromyces bidenticola Hawaii, Mauritius 

MYCOPLASMAS 

aster yellows Hawaii 

Bidens witches broom Brazil 

VIRUSES 

Bidens mosaic 


Sonchus yellow net 

soybean mosaic 

tomato spotted wilt 

Brazil 

Hawaii 

Florida 

Brazil 

Hawaii 

PARASITIC PLANT 

Cassytha filiformis Hawaii 

Table 4.3.3 Species in Agromyzid genera attacking Bidens. 

References 

Linford et al. 1949 

Singh et al. 1979 


Inserra et al. 1989, McSorley 

et al. 198 1 

Rochecouste & Vaughan 1959 

Stevens 1925 

Rochecouste & Vaughan 1959 

Anon 1960, Rochecouste & 

Vaughan 1959 


Vega et al. 198 1 

Kuhn et al. 1982 

Adams 1967 

Christie et al. 1974 

Almeida 1979 

Sakimura 1937 

Raabe 1965 

Genus Specific to Bidens Specific to Coreopsideae Polyphagous 

Melanagromyza 

Amauromyza 

Liriomyza 3 

Calycomyza 

Chromatomyia 

Total 3


Table 4.3.4 Relationship of four major Southeast Asian weeds and some economically 

important genera within the family Asteraceae. 

Family Asteraceae: 21 000 species (Mabberley 1987) 

Tribe Some economically Weed species 

important genera 

Arctoteae 

Carlineae 

Echinopsideae 

Cardueae 

Mutisieae 

Lactuceae 

Vemonieae 

Inuleae 

Astereae 

Eupatorieae 

Calenduleae 

Senecioneae 

Anthemideae 

Heleniae 

Madieae 

Heliantheae 

Tageteae 

Coreopsideae 

Carthamnus, Cynara 

Cichorium, Lactuca 

Aster 

Cineraria 

Chrysanthemum 

Dahlia 

Cosmos, Helianthus, Zinnia 

Ageratum conyzoides, 



Coreopsis Bidens pilosa 

The family Asteraceae, by far the largest in the dicotyledons, has been subdivided into 

18 tribes, some 1300 genera and about 21 000 species (Mabberley 1987). It contains sur- 

prisingly few economically important crop plants, of which lettuce (Lactuca sativa), 

sunflower (Helianthus annuus) and globe artichoke (Cynara scolymus) are the major 

species. However, there are a number of commercially important garden plants, espe- 

cially in the genus Chrysanthemum.



(after Holm etal. 1977)

Map 4.4 Chromolaena odorata 

4.4 Chromolaena odorata 35 


Chromolaena odorata is not a problem weed in the tropical Americas where it evolved. It 

is attacked there by more than 200 insects, at least a quarter of which are probably suffi- 

ciently host specific to be considered as classical biological control agents. The aggres- 

siveness of C. odorata in countries to which it has spread is probably due to the absence 

of most of these natural enemies. 

The arctiid moth Pareuchaetes pseudoinsulata has been established in India, 

Sri Lanka, Philippines, Sabah (Malaysia), the Mariana Is (Guam, Rota, Saipan, Tinian, 

Aguijan) and Federated States of Micronesia (Yap, Pohnpei, Kosrae), but only in the two 

latter island groups has it had spectacular success in controlling the weed. The mite 

Acalitus adoratus has spread naturally to Southeast Asia and Micronesia but, as yet, is 

having minor impact. 

It is probable that a group of natural enemies will be necessary to bring about effec- 

tive biological control of C. odorata in Southeast Asia, but there are a number of species 

that are well worthy of attention and longer term prospects for control appear promising.

36 


4.4 Chromolaena odorata (L.) R.M. King and H. Robinson 

(Formerly Eupatoriurn odoraturn) 

Asteraceae 

Siam weed, devil weed; bizat, tawbizat (Myanmar), tontrem khet (Cambodia), 

French weed (Laos), pokok tjerman (Malaysia), kirinyu, kumpai jepang, rumput 

go1 kar (Indonesia), hagonoy (Philippines) saab sua, yah sua mop (Thailand), co 

hoi (Vietnam) 

Rating 

+++ Msia, Phil 

18 ++ Myan, Thai, Laos, Camb, Viet, Indo 

Origin 

Central America and tropical South America (from Florida to northern Argentina). 

Distribution 

C. odorata is a weed throughout Southeast Asia, Irian Jaya, Papua New Guinea, New 

Britain, Mariana and Caroline Is, southern China, Taiwan, Sri Lanka, Bangladesh, India, 

West, Central and South Africa. 


C. odorata is an upright or scrambling, thicket-forming, perennial shrub, growing from 

1.5 to 3 m high. Its roots are fibrous with a few well formed anchor roots and many later- 

als, the stems round, yellowish, hairy or almost smooth and profusely branched. Its 

leaves are opposite, with toothed margins and are conspicuously three veined. The flow- 

ers are at the tips of all stems, in clusters of 20 to 60, white or pale lilac. The achenes 

consist of 5 mm-long seeds with hooks on their angles, together with a pappus of 5 mm- 

long white bristles. The leaves have a pungent odour when damaged. Seed production is 

prolific (as many as 2 million per plant) and seeds provide the main mode of reproduc- 

tion. The achenes float long distances in the air and the seed hooks cling to hair and 

clothing. Germination occurs as soon as there is adequate moisture, although some 66% 

of seeds are not viable. Buried seeds lose up to 50% of their viability after 2 years (Yadav 

and Tripathi 1982). 

Importance 

C. odorata is not a serious weed in the Americas and no specific control methods are 

necessary (McFadyen 1991a). This is in stark contrast to its serious weed status in the 

countries to which it has spread and has been attributed to the many natural enemies that 

attack it in the Americas (McFadyen 1989, 1991~). It was introduced to Calcutta in the 

1840s, had spread into Sri Lanka, Southeast Asia and Nigeria by the 1940s and into Irian 

Jaya, New Britain and Micronesia by the 1980s. It is forecast to spread widely and


aggressively in equatorial Africa, northern and eastern Australia and the Pacific 

(McFadyen 1988b, 1989). 

C. odorata grows in many soil types, but prefers well drained conditions and an 

annual rainfall above 1000 mm. Although it is not a problem in continuously cultivated 

land, it is most common and causes most losses in plantation crops, including coconut, 

rubber, oil palm, tea, coffee, cocoa, teak and cashew. It also thrives in areas newly 

cleared for planting, in abandoned or neglected fields, wastelands and along roadsides. It 

is sometimes a weed in pastures. Its rapid growth enables it to smother most competitors 

and it inhibits many with its allelopathic properties. It dies back after flowering in areas 

with a pronounced dry season and then becomes a fire hazard. After burning or cutting, 

the plants shoot freely from the crown. They are capable of forming dense tangled bushes 

two to three metres high, occasionally reaching six metres as climbers on other plants. 

The stems branch freely, with 20 or more laterals developing from axillary buds and 

often bent over under their own weight. Impenetrable stands of the weed cut off access to 

pastures and provide hiding places for rats, pigs and other undesirable animals. C. 

odorata is intolerant of shade, so that it dies out when the canopy closes in plantations 

(Ambika and Jayachandra 1990, McFadyen 1988b, 199 1 a). The shoots and young leaves 

contain nitrate at levels 5 to 6 times those toxic to stock and also pyrrolizidine alkaloids 

and cattle deaths occur following grazing. Hand weeding of Chromolaena is reported to 

cause skin allergy and scratches to result in infections (Ambika and Jayachandra 1990). 

It is interesting that the spread of C. odorata in West Africa has led to a 

polyphagous grasshopper Zonocerus variegatus becoming a pest. Although they are 

unable to mature on the weed as the only diet, hoppers are strongly attracted to the plant 

and especially to its flowers; and thickets are preferred night roosting sites. BopprC 

(1 991) hypothesises that the pyrrolizidine alkaloids accumulated from feeding on C. 

odorata protect the grasshoppers and their eggs from predators and parasitoids, leading 

to increased fitness and population density. However, this only occurs during the dry season, 

but not in the wet season when C. odorata does not bloom. 

Claims have been made (e.g. Field 1991, Herren-Gemmill 1991) that, under some 

circumstances, C. odorata may be beneficial to resource-poor farmers. One potential 

advantage, is its ability to outcompete another serious weed, alang-alang (Imperata cylindric~). 

However, McFadyen (1992) pointed out that a suitable perennial legume would 

be even more beneficial than C. odorata, and she also refuted a number of other claims. 

In Sri Lanka the indigenous legume Tephrosia purpurea has been successfully used to 

suppress weeds including C. odorata under coconut (Salgado 1972). Whatever potential 

benefits there may be in the presence of C. odorata there is an enormous body of fact to 

demonstrate that C. odorata has serious adverse effects on agricultural productivity in 

countries to which it has been introduced. 


A good deal is known about the insects attacking Chromolaena odorata, mainly as a 

result of studies aimed at biological control which started in the late sixties at the 

Commonwealth Institute of Biological Control Station in Trinidad. A number of scientists

38 


were involved, but principally R.E. McFadyen (nee Cruttwell) (Bennett and Cruttwell 

1973, Bennett and Rao 1968, Bennett and Yaseen 1975, Cock 1984a, Cock and Holloway 

1982, Cruttwell 1973a,b, 1974, 1977a,b, Cruttwell and Bennett 1969, McFadyen 

1988a,b, 199 1 a,b, Yaseen and Bennett 1977). 

An extensive bibliography dealing with all aspects of C. odorata, including its natural 

enemies and biological control, was compiled by Muniappan et al. (1988a), later 

supplemented in Chromolaena odorata Newsletters 3 (1990) and 6 (1992). The proceedings 

of three International Workshops on Biological Control of Chromolaena odorata, 

held in 1988, 1991 and 1993 also contain a wealth of up-to-date information. 

In the Americas C. odorata is attacked by at least 207 insect and 2 mite species 

(McFadyen 1988a). Of these, about half are probably polyphagous, a quarter are restricted 

to the Asteraceae and a quarter specific to Chromolaena. All stages of growth of the 

above ground parts of the plants are attacked, but the roots have not been examined 

(McFadyen 1991a) and not all regions where C. odorata occurs naturally were visited. 

For other regions of the world McFadyen (1988a) quotes records of 42 insect and 9 mite 

species, the vast majority of which are, or are likely to prove, polyphagous. Since then a 

few additional species have been recorded, all but one of which (an eriophyid mite, see 

India below) are likely to be polyphagous. 

In Trinidad, the cumulative effect of the natural enemies is great, between 25 and 

50% of all growing tips being destroyed. Seed germination is as low as 17% and many 

flowerheads fail to produce seed. Seedlings often succumb to the attack of stem and tip 

feeding insects and competitiveness and growth of established plants is greatly reduced 

by insect attack. At different sites and in different seasons damage is caused by different 

insects and, in general, is heaviest in shaded sites. Some of the insects are heavily 

attacked by parasitoids and if introduced without these to another country might prove to 

be even more effective. 

In addition to an arctiid moth (Pareuchaetes pseudoinsulata) and a weevil (Apion 

brunneonigrum), which have already had considerable attention paid to them (see next 

section), McFadyen (1 99 1 c) has nominated an additional 1 1 insects for priority evaluation 

(Table 4.4.1). Furthermore, others (Cruttwell 1974, Cock 1984a, Cock and Holloway 

1982, McFadyen 1988c, Muniappan and Viraktamath 1986) have suggested an additional 

22 species (Table 4.4.2) which were evidently considered less important by McFadyen 

(1991~). It is clear therefore that, if required, there are many promising candidates for 

detailed consideration. The additional species of Pareuchaetes suggested by Cock and 

Holloway (1982) have not been investigated in detail, but all are believed to breed on C. 

odorata or related species and several may be better adapted climatically and biologically 

than P. pseudoinsulata to conditions in many overseas countries. 

Although no special search has been carried out except in Trinidad and Tobago a 

number of fungal pathogens occurring on C. odorata are shown in table 4.4.3. Half of the 

records 'come from outside its area of origin and must, therefore, be suspected of having a 

wider than desirable host range. Possibly Cionothrix praelonga is of greatest interest, 

since preliminary tests indicate that it may be host specific (Ooi et al. 1991). It is autoecious 

(i.e. it does not have an alternate host), occurs in the Caribbean and Venezuela and


Table 4.4.1 Potential biological control agents for C. odorata: insects (after 

McFadyen 199 1 c). 

Species Part Damage Problem Country found 

attacked 

Coleoptera 

CHRYSOMELIDAE 

Aulacochlamys sp. 

Chlamisus insularis 

Pentispa explanata 

CURCULIONIDAE 

Rhodobaenus 

cariniventris 

Diptera 

AGROMYZIDAE 

Melanagromyza 

eupatoriella 

CEClDOMYllDAE 

Clinodiplosis sp. 

Perasphondylia 

reticulata 

TEPHRlTlDAE 

Procecidochares sp. 

Lepidoptera 

BUCCULATRICIDAE 

Bucculatrix sp. 

NYMPHALIDAE 

Actinote anteas 

PYRALIDAE 

Mescinia parvula 

stem 

stem 

leaf miner 

stem 

shoot borer 

shoot galls 

bud galls 

stem galls 

leaf miner 

leaf 

shoot borer 

moderate 

minor 

moderate 

great 

great 

great 

great 

moderate 

minor 

great 

great 

Trinidad 

all Americas 

prefers shade Trinidad 

Trinidad 

cage mating West Indies, S. America 

rearing Trinidad 

cage rearing all Americas 

parasites Americas 

Mexico 

cage mating Trinidad, Costa Rica 

cage mating all Americas 

causes conspicuous leaf lesions. Pseudocercospora eupatorii-formosani is reported to be 

common and damaging on C. odorata in Brunei, but is widespread already in South and 

Southeast Asia (Chacko 1988, Evans 1987, Peregrine and Ahmad 1982). 


Four insects have been released for biological control, the weevil Apion brunneoni- 

grurn, the fly Melanagromyza eupatoriella and two moths Mescinia parvula and 

Pareuchaetes pseudoinsulata (Table 4.4.4). Of these, only the last has become estab- 

lished, fairly readily in Sri Lanka, Guam and other Micronesian islands, but with some 

difficulty in India and Sabah (Malaysia) and it 'has since spread unaided to the 

Philippines and Brunei. It failed to become established in Thailand, Ghana, Nigeria and 

South Africa. It has produced spectacular defoliation and death of many plants in Guam


Table 4.4.2 Additional potential biological control agents for C. odorata: arthropods. 

Species Part attacked Country found 

Coleoptera 

CERAMBYCIDAE 

Aerenica hirticornis stem borer Trinidad, Bolivia, 

Brazil, Argentina 

CURCULIONIDAE 

Baris sp. 

Centrinaspis sp. 

Diptera 

CEClDOMYllDAE 

Asphondylia corbulae 

Clinodiplosis eupatorii 

Clinodiplosis sp. 

Contarinia sp. 

Neolasioptera brickelliae 

Neolasioptera cruttwellae 

Neolasioptera eupatorii 

Neolasioptera frugivora 

TEPHRITIDAE 

Cecidochares jluminensis 

Procecidochares connexa 

flowers, leaf buds 

flowers, leaf buds 

flower galls 

leaf galls 

bud galls 

flowers (achenes) 


stem galls 

stem galls 


flowers 

stem galls 

Trinidad 

Trinidad, Costa Rica 

El Salvador, Trinidad 

Central America, Brazil, West 

Indies 

Trinidad 

Trinidad 

Trinidad 

Trinidad 

USA, Trinidad, Bolivia 

Trinidad 

Trinidad, SE Brazil 

Mexico, Brazil, Bolivia 

Lepidoptera 

ARCTllDAE 

Pareuchaetes aurata aurata leaves. buds Paraguay, Argentina, 

Pareuchaetes aurata aurantior 

Pareuchaetes arravaca 

Pareuchaetes insulata 

Pareuchaetes misantlensis 

Pareuchaetes sp. 

GELECHllDAE 

Dichomeris (= Trichotaphe) sp. 

leaves, buds 

leaves, buds 

leaves, buds 

leaves, buds 

leaves, buds 

leaf roller (see 

Cruttwell 1973b) Trinidad 

SE Brazil, Bolivia 

Amazon River 

Surinam and French Guiana 

Southern USA, Mexico, 

Caribbean, Colombia 

Mexico 

Mexico 

LYCAENIDAE 

Calephelis laverna leaves Brazil, Trinidad, Venezuela, 

Central America 

Acarina 

ERlOPHYlDAE 

Calacarus sp. shoots 

India

Table 4.4.3 Natural enemies of C. odorata: fungi. 


Species Country found References 

Anhellia niger 

Cercospora sp. 

Cercospora eupatorii 

Cercospora eupatoriicola 

Cercospora eupatorii-odoratii 

Cionothrix praelonga 

Fusarium oxysporum f. sp. elaeidis 

Guignardia eupatorii 

Mycovellosiella perfoliata 

Phoma sp. 

Phomopsis eupatoriicola 

Phyllosticta eupatoriicola 

Pseudocercospora eupatorii-formosani 

Septoria sp. 

Septoria ekmaniana 

Trinidad, Tobago 

Peninsular Malaysia, 

Sabah 

North America, Cuba, 

Nepal, India, Ivory Coast 

India, Bangladesh 

Malaysia 

Dominica, Tobago, 

Venezuela 


Sri Lanka 



not recorded 

not recorded 

India, Myanmar, Thailand, 

Malaysia, Borneo, Brunei, 

Guam 


Ooi et al. 1991 

Singh 1980 

Chacko 1988, Evans 1987 




Oritsejafor 1986 







Russo 1985 


and striking but sporadic defoliation in Sri Lanka. In India, populations have built up but 

damage has seldom been great. Where established, it is heavily attacked by a range of 

predators and these are believed to have prevented successful establishment in several 

countries. 

The eriophyid mite Acalitus adoratus causes abnormal growth of the epidermal 

hairs on young leaves and stems of C. odorata. Although it was never purposely intro- 

duced, it was observed in Thailand in 1984 and the Philippines in 1987, but had probably 

been present for some years. It is also widespread in Java and Sumatra, but there is no 

information from other Indonesian islands. It is present in Yap and Palau in the Caroline 

Islands and was observed on Guam in November 1993 (R. Muniappan pers. comm.). It is 

not present in India and it is not known whether it is present in Sri Lanka (Cruttwell 

1977b, McFadyen 1991 b, 1993, Muniappan et al. 1988a). 

Further details follow of the situation in individual countries and of the biology of 

some of the more promising natural enemies. 

Asia 

INDIA 

In one study, 11 insects and 3 mites were found attacking C. odorata. All except 

Calacarus sp. (Eriophyidae) are polyphagous (Muniappan and Viraktamath 1986, 

Viraktamath and Muniappan 1992). Most eriophyid mites have a highly restricted host 

range, so it is not clear whether it may even have accompanied the weed from the


4.4.4 Introductions for the biological control of Chromolaena odorata. 

Species Country Liberated Result References 

Coleoptera 

BRENTHIDAE 

Apion brunneonigrum Ghana 1975 - Cock 1984a, 1985, 

Guam 1984 - Nafus & Schreiner 1989 

India 1972-83 - Chacko & Narasimham 

1988, Cock 1984, 1985 

Malaysia 1970 - Ooi et al. 1988a,b 

(Sabah) 

Nigeria 197CL75 - Cock 1984a, 1985 

Sri Lanka 197676 - Cock 1984a, 1985 

Lepidoptera 

ARCTllDAE 

Pareuchaetes aurata 

aurata 

Pareuchaetes 


South Africa 

Pohnpei 

Ghana 

Guam 

India 

Indonesia 

Kosrae 

Malaysia 

(Sabah) 

Northern 

Marianas 

Nigeria 

South Africa 

Sri Lanka 

Thailand 

Yap 

Julien 1992, 

Kluge & Caldwell 1993 

Esguerra et al. 199 1, 

Esguerra et al. 1994 

Muniappan et al. 1988b 

Cock 1985, 

Cock & Holloway 1982 

Julien 1992 

Nafus & Schreiner 1989, 

Seibert 1989 

Chacko & Narasimham 

1988, Cock & Holloway 

1982 

Chacko & Narasimham 

1988, Julien 1992, 

Muniappan et al. 1989, 

Satheesan et al. 1987 

McFadyen pers. comm. 

Esguerra et al. 1994 

Cock & Holloway 1982, 

Ooi et al. 1988a,b, 

Syed 1979a 

Nafus & Schreiner 1989, 

Seibert 1989 

Cock & Holloway 1982 

Julien 1992, Kluge 1991, 

Kluge & Caldwell 199 1 

Dharmadhikari et al. 1977 

Napompeth et al. 1988 

Marutani & Muniappan 

199 1 a, Muniappan et al. 

1988b 

PY RALl DAE 

Mescinia parvula Guam Nafus & Schreiner 1989


Americas. In another study 21 polyphagous insects were recorded from C. odorata, of 

which the most widespread and numerous were Aphis fabae and A. spiraecola (Lyla and 

Joy 1992, Lyla et al. 1987). Some of these same species are included amongst the 31 

insects and 9 mites recorded on Chromolaena by Chacko and Narasimham (1988). 

Pareuchaetes pseudoinsulata from Trinidad was cleared of a nuclear polyhedral 

virus and mass reared. It was first released in 1973 at several sites in Karnataka, but no 

establishment occurred. Observations suggested that two ants, Myrmicaria brunnea and 

Oecophylla smaragdina, were major predators (Cock and Holloway 1972). P. pseudoin- 

sulata from Sri Lanka (where it had been sent and had already become established) was 

next released and appeared to be doing well until unexpectedly wiped out by virus (Cock 

1985). However, further material from Sri Lanka was laboratory reared and 36000 larvae 

and 1000 adults released from 1984 onwards, this time in Kerala. This procedure resulted 

in field establishment (Chacko and Narasimham 1988, Joy et al. 1993, Muniappan et al. 

1989, Satheesan et al. 1987). Most recently, the establishment of P. pseudoinsulata at 

Sullia Taluk in Karnataka State and defoliation of Chromolaena thickets over about 1000 

km2 was reported in December 1992 (R. Muniappan pers. comm. 1993). However the 

overall performance of the moth has been unsatisfactory (Joy et al. 1993). 

Small releases of the weevil Apion brunneonigrum have been made since 1972, but 

establishment has not resulted (Cock 1985, Ooi et al. 1991). 

SRI LANKA 

R pseudoinsulata was received from India in 1973 and about 2000 larvae released in a 

coconut estate in the North Western Province. Six months after release spectacular defo- 

liation was observed of a hectare of previously impenetrable growth of C. odorata. In 

addition to leaves, terminal buds and tender stems were being consumed. Further releas- 

es were made and two years later it was estimated that some 800 ha of C. odorata had 

been defoliated (Dharmadhikari et al. 1977). Since then sporadic, heavy defoliation has 

mainly occurred at the beginning of the dry season at the time of flowering. This has 

caused great damage and, at times, death of the weed. However R pseudoinsulata popu- 

lations fluctuate considerably, due in no small measure to natural enemies. Young larvae 

are taken by birds and predatory Sycanus bugs. They are also parasitised by the braconid 

Apanteles creatonoti and the tachinid Exorista sp. (Kanagaratnam 1976). In one series 

of experiments from 63 to 100% of pupae were consumed by ants, termites and lizards 

(Mahindapala et al. (1980). Perera (1981) fed P. pseudoinsulata larvae on C. odorata 

leaves dipped in 32P labelled sodium orthophosphate, transferred them to C. odorata in 

the field and collected predators from pitfall traps. Several carabids and a histerid 

showed no radioactivity, nor did the ants Odontomachus simillimus and Diacamma 

rugosum which were observed carrying away treated larvae to their nests, indicating that 

they do not feed on the larvae soon after capture. There was no unusual preponderance 

of predatory wasps, but birds were observed picking up larvae so it is likely that they 

were the cause of the sudden decline in larval population (P.A.C.R. Perera pers. comm. 

1993).


How effective the moth is as a control agent is yet to be determined. However, 

Perera (1981) calculated that a t? pseudoinsulata larva from hatching to pupation con- 

sumes an average of 184.6 cm2 of leaf. Based on measurements of a heavy growth of C. 

odorata, there are 22.42x lo8 cm2 of leaf area per ha requiring about 12 million larvae to 

produce defoliation. Assuming an average egg production of 200 per female moth and a 

1 : 1 sex ratio, 12 million larvae could be produced in two generations (3 months) with a 

release of 600 to 700 females. Cock and Holloway (1982) have suggested that there is a 

better climate match between Sri Lanka and Trinidad than for most of the other countries 

where the moth has been released. 

Apion brunneonigrum were released between 1974 and 1976 and, two months after 

release, were seen on flower heads but have not been recovered since (Cock 1985, 

Kanagaratnam 1976, Ooi et al. 1991). 

Southeast Asia 

BRUNEI 

Although no releases of P. pseudoinsulata have been made, two females were trapped in 

the early 1980s, presumably having resulted from the colonies established in neighbour- 

ing Sabah (Malaysia) (Cock 1985). 

INDONESIA 

An aphid has been observed to attack young shoots and cause leaf curl of C. odorata. 

Work on biological control of the weed was initiated in 1991 with the introduction of P. 

pseudoinsulata to Sumatra, but there is no information on the outcome. There is a current 

project (1993) under R.E. McFadyen to study the host specificity of the tephritid fly 

Procecidochares connexa and either the moth Mescinia parvula, the stem boring 

Melanagromyza eupatoriella or the butterfly Actinote anteas with a view, if judged safe 

to do so, to liberation in Indonesia and the Philippines (R.E. McFadyen pers. comm., 

Tjitrosoedirdjo 1991, Tjitrosoedirdjo et al. 199 1). 

MALAYSIA (SABAH) 

Aphis spiraecola attacks young shoots of C. odorata and causes leaf fall (Bennett and 

Rao 1968). 

t? pseudoinsulata was introduced from India to Sabah in 1970 and releases made 

between 1970 and 1974 of over 4000 eggs, 40 000 larvae and 700 adults. Temporary 

establishment occurred in two areas in 1973 and 1974, but both colonies then appeared to 

die out over the next couple of years. This was considered to be due to general predators, 

such as ants (Cock and Holloway 1982). However, in 1983 and 1987, pockets of larvae 

appeared scattered over Sabah and often distant from the sites of original release (Ooi et 

al. 1988a,b). There is a good climate match between Sabah and Trinidad, which may 

explain the establishment (Cock and Holloway 1982). 

Small releases of A. brunneonigrum were made in 1970 and recoveries were report- 

ed a year later, but there is no indication that the weevil has survived (Ooi et al. 1988a,b, 

Syed 1973, 1975, 1979a,b).


PHILIPPINES 

Aphis gossypii, A. spiraecola (= A. citricola) and the tortricid Homona coflearia were 

found attacking C. odorata (Torres 1986) and the total of natural enemies increased to 11 

by 8 additional (unspecified) insects (Torres 1988). Although it had not been intentional- 

ly introduced, numerous larvae of P. pseudoinsulata were discovered in 1985 in a limited 

area near the coast of Palawan. They were feeding on the leaves and stems of C. odorata 

under coconut trees and along roads, but surveys elsewhere at the time revealed no evi- 

dence of P. pseudoinsulata larvae (Aterrado 1986a,b, Torres and Paller 1989). However 

P. pseudoinsulata was discovered later in Zamboanga City, Bohol and northern Leyte 

provinces in the Visayas islands (Aterrado and Talatala-Sanico 1988). 

The eriophyid mite Acalitus adoratus was discovered in the Philippines in 1987 

(McFadyen 1 99 1 b). 

THAILAND 

A number of insects were found attacking C. odorata: the aphids Aphis craccivora, 

A. gossypii and A. spiraecola, the weevil Hypolixus trunculatus, a stem boring cicindelid 

larva and the arctiid moth Amsacta lactinea. They were causing little damage 

(Napompeth 1990a,b, Napompeth et al. 1988, Napompeth and Winotai 199 1). 

P. pseudoinsulata was introduced from Guam from 1986 to 1988 but, despite 

repeated field releases in 1987 and 1988, did not become established. The shoot miner 

Melanagromyza eupatoriella was introduced from Trinidad in 1978, but could not be 

reared and was not released. 

The mite Acalitus adoratus, detected in 1984, has since spread to all C. odorata 

infested areas, but is not having a significant effect on the weed. 

VIETNAM 

Infestations of Aphis craccivora and A. gossypii have been recorded on C. odorata, but 

no releases of biological control agents have been made (Napompeth and Hai 1988). 

Pacific 

GUAM 

P. pseudoinsulata was introduced from India and Trinidad, mass reared and first released 

in Guam in 1984 and later in the Northern Marianas. Initially late instar larvae were 

released in batches of up to 800, but were heavily attacked by ants, spiders, toads and 

other general predators and failed to become established. Next, groups of 500 or more 

adult moths were released at a number of sites, resulting in establishment in all release 

areas. Populations built up rapidly, defoliation of Chromolaena soon followed and almost 

all plants were stripped. Shoots arising from the crowns were also attacked as they 

appeared and, within a year, over 90% of the plants were killed. The moth spread rapidly 

and by 1987 had reached almost all infested areas of Guam. Eventually more than 

25 000 ha of the weed had been defoliated (Muniappan 1988c, Nafus and Schreiner 1989, 

Seibert 1989). A parasitoid Exorista xanthaspa (= E. civiloides) caused up to 30% mor- 

tality and predation by ants, spiders, toads and lizards occurred (Seibert 1989).


It was observed that the feeding of P. pseudoinsulata larvae caused the leaves of C. 

odorata to turn yellow, an effect that could not be produced by simply applying larval 

excreta to the plant. Yellow leaves were tougher and had a higher level of nitrate and, 

when larvae were forced, much against their preference, to feed on yellow leaves, they 

exhibited slow growth and high mortality. Furthermore, larvae continued to feed on yel- 

low plants both by day and night (exposing them to daytime predators), whereas on green 

plants they fed at night and hid at ground level by day (Marutani and Muniappan 1991 b). 

Interestingly, the yellow plants appear to lose their allelopathic properties and hence this 

major aid to dominance over other vegetation. The yellowing is reversible if the insects 

are removed (McConnell et al. 1992, Muniappan and Marutani 1992). 

Three additional insects were released to aid in the control of Chromolaena, but 

there is no evidence of establishment. Apion brunneonigrum was released early in 1984 

at the beginning of the dry season when the above ground growth of Chromolaena dies 

back. Because of the unsuitable condition of the host plants the beetle was not expected 

to become established. Small numbers of Mescinia parvula were released late in 1984 

and again late in 1986 (Seibert 1989). The mite Acalitus adoratus appeared in Guam in 

1993 (R. Muniappan pers. comm.). 

Larvae of the pyralid moth Eucampyla etheiella were observed attacking young 

flower buds and mature flowers and causing extensive damage. Larvae were parasitised 

by the eulophid Elachertus sp. and the elasmid Elasmus sp. (Marutani and Muniappan 

1 990). 

NORTHERN MARIANAS (ROTA, TINIAN, SAIPAN, AGUIJAN) 

P. pseudoinsulata has been established from liberations in 1986 and 1987 on all of these 

islands (Muniappan et al. 1989, Nafus and Schreiner 1989). 

Federated States of Micronesia 

KOSRAE 

Monthly releases of P. pseudoinsulata larvae in batches of 1000 to 4000 were made from 

early 1992 in sunny areas and defoliation of C. odorata was observed six months later. 

Predators were less active in sunny than in shady locations (Esguerra et al. 1994). 

PA LA U 

Although no releases of biological control agents have been made, the mite Acalitus ado- 

ratus was found to be present (Muniappan et al. 1988b). 

POHNPEI 

P. pseudoinsulata larvae were introduced from Guam in 1988, some liberated and others 

mass reared during which both larvae and adults were released until 1992. In four release 

sites extensive feeding injury and heavy defoliation of C. odorata was observed in 1991 

and populations persisted in 1992 in burnt areas where Siam weed was regenerating from 

root stocks (Esguerra et al. 1994). Heavy predation, especially in shaded conditions, was 

observed on all stages by ants, spiders, birds and lizards (Esguerra et al. 1991, 1994).


YAP 

C. odorata was first reported in 1987. P. pseudoinsulata was released in 1988 at 14 dif- 

ferent sites, but failed to become established except at one location where only 100 lar- 

vae and 104 adults had been released (Muniappan et al. 1988b). It eventually disappeared 

at this site. However releases of 500 larvae in September and October and several hun- 

dred in December 1990 to June 1991 resulted in establishment (Marutani and Muniappan 

1991a). As on Guam and Rota, larvae of Eucampyla etheiella were found causing exten- 

sive damage to buds and mature flowers (Marutani and Muniappan 1990). The eriophyid 

mite Acalitus adoratus was found attacking C. odorata late in 1988, although it was not 

observed during a survey of the weed in May of that year (Muniappan et al. 1988b). 

Africa 

GHANA 

P. pseudoinsulata from India was used to establish a culture and releases were made 

between 1973 and 1978 in a variety of habitats including oil palm plantations. Although 

small amounts of leaf damage were observed shortly after release, no recoveries were 

made. Failure to establish was ascribed to predators, in particular to ants (Cock and 

Holloway 1982). 

One small release of Apion brunneonigrum was made in 1975, but it failed to 

become established (Cock 1985). 

NIGERIA 

P. pseudoinsulata shipped from Ghana between 1973 and 1978 were released, but no 

establishment occurred (Cock and Holloway 1982). 

A. brunneonigrum was sent from Trinidad from 1970 to 1975, but there is no record 

of establishment (Cock 1985). 

SOUTH AFRICA 

Disease-free adults of P. pseudoinsulata originating from Guam were released in batches 

of 500 to 1000 at 10 sites in Natal in 1989, but there are no signs of establishment. Very 

heavy egg predation (up to 82%) by ants and chrysopids was observed (Kluge 1991, 

Kluge and Caldwell 1991). P. pseudoinsulata has been obtained from Florida where the 

climate is similar to that in Natal and where there is a rich ant fauna. It is (as of 1991) to 

be released as soon as laboratory cultures of larvae have been cleared of microsporida. 

The larvae of another arctiid moth, Pareuchaetes aurata aurata, from Chromolaena 

jujuensis in Argentina were found to feed voraciously and complete their development on 

C. odorata. Females scatter their eggs around the base of the host plant and it is hoped 

that this will help to overcome the problem of ant predation. After specificity testing it 

has been released in Natal, but no further information is available (Kluge and Caldwell 

1993). 

'A laboratory culture of the butterfly Actinote anteas has been established with 

material collected in Costa Rica and host testing is to commence (Kluge and Caldwell 

1991). 

Work is also in progress on the host specificity of the weevil Rhodobaenus


cariniventris and a leaf spot disease caused by Septoria sp. (Kluge and Morris 1992). 

Major natural enemies 

Acalitus adoratus Acarina: Eriophyidae 

Recorded originally from Trinidad, Florida, Brazil and Bolivia, it appeared without spe- 

cial assistance in Thailand, Philippines, Indonesia (Java and Sumatra), Caroline Is (Yap, 

Palau) Guam and southern China. It was not present in India in the mid 1980s nor in Sri 

Lanka or West Africa (McFadyen 1993). 

These tiny mites (0.1 4 to 0.1 8 mm long) usually live on the lower surface of leaves. 

Their feeding induces abnormal growth of the epidermal hairs, resulting in the formation 

of erineum patches, the term given to areas covered with dense twisted hairs amongst 

which the mites live. These appear as white patches on the leaves, usually 0.5 to 3 mm in 

diameter, and the whole leaf surface may be affected. The patches often turn yellow on 

older leaves. The nymphs and adults feed, and the eggs are laid, between the epidermal 

hairs. Particularly heavy infestations develop in dry and exposed situations and, although 

the damage is not spectacular, heavy attack stunts, distorts and slows growth, thereby 

reducing competitiveness. Tests indicated that, as with many other eriophyid mites, 

A. adoratus is host specific. 

When infested leaves senesce, A. adoratus leave the erineum patches and are preyed 

upon by other mites and by the larvae of a cecidomyiid fly, Arthrocnodax meridionalis 

(Cruttwell 1977b, McFadyen 199 1 b, Muniappan et al. 1988a,b). 

It is suggested that A. adoratus was accidentally introduced to Sabah when field- 

collected adults of the weevil Apion brunneonigrum in Trinidad were released directly in 

the 1970s; and that it has since spread naturally and on leaves of C. odorata used as 

packing material around fruit and other produce (McFadyen 1993). 

Actinote anteas Lepidoptera: Nymphalidae 

Recorded from Costa Rica and Trinidad. The host specificity of this acraeinine defoliator 

is being examined in South Africa (Kluge and Caldwell 1991). 

Apion brunneonigrum Coleoptera: Apionidae 

This weevil has been recorded from Trinidad, Venezuela and Argentina. Small releases 

were made in West Africa, India, Sri Lanka, Sabah and the Marianas, but the weevil per- 

sisted only in Sabah and then apparently only briefly. The reasons for these failures have 

not been investigated. 

Cruttwell (1973a) studied its biology and host specificity and found that it would 

feed and develop only on C. odorata and C. ivaefolia. It has never been recorded damag- 

ing economic plants either in Trinidad or South America. The life history is closely linked 

with the development of its host, the adults becoming reproductively mature at the time 

that the plant produces young flower buds which provide food for egg maturation. Eggs 

are deposited in the developing flower heads and larvae feed within the flower heads, 

destroying the seeds. Pupation occurs in the flowerheads and, until the next flowering, 

adults feed on tender growth, usually in shaded situations, and may do considerable dam-


age (Cock 1984a). An individual larva destroys 30 to 60 seeds during development and 

the ovipositing female many young flowers. A. brunneonigrum thus has potential for 

causing considerable damage, particularly in lightly shaded conditions. 

Aulacochlamys sp. Coleoptera: Chrysomelidae 

Widespread and occasionally abundant in Trinidad, where it causes moderate damage. 

Eggs are laid singly in a cylindrical ribbed case formed from faeces. These cases form 

the apex of a conical larval case which is enlarged as the larva grows. Larvae feed on the 

surface of stems and leaf petioles. Mature larvae attach the 3.5 to 3.7 mm long case to a 

stem and pupate inside. Adults emerge one to two weeks later. The small black adults 

(1.8 to 2.5 mm long) feed on the surface of stems and petioles. No parasitoids are known 

(McFadyen 1988a). 

Bucculatrix sp. Lepidoptera: Bucculatricidae 

Recorded from C. odorata in Mexico and Chromolaena jujuensis (= Eupatorium hooke- 

rianum) in Argentina. Larvae are solitary leaf miners and pupate in the mines (McFadyen 

1988a). 

Chlamisus insularis Coleoptera: Chrysomelidae 

Recorded from Mexico, Panama and Trinidad. The life history is similar to that of 

Aulacochlamys sp., but this species is somewhat larger. The mature larval case is conical 

with a rough surface and 6 to 7 mm long. The adults are black with golden markings and 

3.3 to 4.3 mm long. Adults are known to feed on C. odorata, C. ivaefolia and Bidens 

pilosa. A black, solitary eulophid egg parasitoid is known (McFadyen 1988a). 

Clinodiplosis sp. Diptera: Cecidomyiidae 

Recorded from C. odorata and C. ivaefolia in Trinidad. Up to three larvae at a time live 

between the bud leaves of stem tips or axillary buds, destroying tissue and preventing 

growth. A gall is formed by the slight swelling of the bud leaves which become red and 

densely covered with hairs. Mature larvae drop to the ground and pupate just below the 

soil surface. Adults emerge 11 to 18 days later. This gall midge is abundant and wide- 

spread in Trinidad, breeds throughout the year and causes considerable damage to C. 

odorata (McFadyen 1988a). 

Mescinia parvula Lepidoptera: Pyralidae 

Recorded from Trinidad; similar larvae were found on C. odorata in Mexico and Brazil 

and on C. jujuensis in Argentina. A few individuals were released on Guam in 1984, but 

there has been no evidence of establishment. 

Ovipositing females select leaves with dense hairs (in effect young leaves) with the 

result that developing buds are nearby. Eggs are laid individually amongst the epidermal 

hairs on the underside of the young leaves and hatch in 5 to 6 days. Young larvae move 

to a terminal or axillary bud and several may enter the same bud. They bore down the 

stem destroying meristematic tissue and preventing growth. Larvae may leave a stem and 

enter a new bud. After 13 or so days larvae leave the stem to spin a flimsy cocoon, either 

attached to the plant or among ground litter, in which they pupate. Adults emerge 10 to


11 days later and live up to 6 days. Attempts to induce mating in cages in Trinidad were 

unsuccessful. In specificity tests larvae fed on only a few Asteraceae other than C. odora- 

ta and C. ivaefolia but, with the exception of 1 out of 30 larvae placed on Dahlia, no 

development was ever completed. Over a three year period in Trinidad, no oviposition or 

attack was observed on Dahlia plants growing near C. odorata which was frequently 

attacked by M. parvula. Furthermore, since Dahlia leaves are not hairy, it is most unlike- 

ly that M. parvula would ever oviposit on them. Larvae in Trinidad are attacked by eight 

hymenopterous and one tachinid parasitoid and, if freed from these, might do consider- 

ably more damage to C. odorata (Cruttwell 1977a). 

Pareuchaetes aurata aurata Lepidoptera: Arctiidae 

This subspecies occurs in south-eastem Brazil, Paraguay and northern Argentina at lati- 

tudes (26" to 30°S) similar to those of Natal, South Africa. It has an average life cycle of 

30 days at 26' to 29OC and 58 days at 22O to 25OC. Its larvae are nocturnal feeders and 

shelter at the base of plants during the day. In the laboratory, pupation occurred in a flimsy 

cocoon spun between leaves on the plant. Mating may occur on the night of emergence 

and an average of 242 eggs are laid over the next eight days. These are laid singly on the 

ground and, in the laboratory, newly-hatched first instar larvae were able to walk up about 

2m of stem to commence feeding. In the field ?! aurata aurata is found in shaded habitats 

near surface water. 

In Argentina, larvae and pupae are infected with a microsporidan disease (Nosema 

sp.), up to 20% of larvae are parasitised by a complex of braconid, chalcidid and tachinid 

parasitoids and all stages are subject to attack by predatory ants. 

The usual host plant of ?! aurata aurata is Chromolaena jujuensis, but it has been 

successfully reared for more than 10 generations on C. odorata. In the field it has never 

been recorded as a pest on any of the many commercially important crops grown in its 

natural area of distribution. It was liberated in Natal (South Africa) in 1990 (Kluge and 

Caldwell 1993). 

Pareuchaetes pseudoinsuhta Lepidoptera: Arctiidae 

This moth is native to Trinidad, Tobago and the north-eastern coast of Venezuela. It has 

become established in Brunei, Guam, India, Philippines, Sabah, Sri Lanka, the Northern 

Marianas and Yap, but has failed to do so in Ghana, Nigeria, South Africa and Thailand. It 

was previously misidentified first as Ammalo arravaca and then as A. insulata, which is a 

closely related but distinct species (Cock and Holloway 1982). 

The moth, which lives up to about 10 days, lays 150 to 250 eggs (maximum 580) in 

groups attached to the lower surface of the leaves of C. odorata. Larvae feed on the leaves 

and are gregarious until the 3rd instar, but then disperse. From the 4th instar on they feed 

at night, hiding by day amongst debris at the base of the plant, where they later pupate. 

The life cycle varies from 40 to 60 days and breeding occurs throughout the year. Host 

specificity studies in Trinidad showed that development occurred only on Chromolaena 

ivaefolium, C. microstemon and C. odoratum. In addition larvae developed, but only as far 

as the 3rd instar, on Ageratum conyzoides (Bennett and Cruttwell 1973), although in Sri 

Lanka, adults were produced on this weed. However their eggs had a somewhat lower


hatchability than those from adults bred on C. odorata (Mahindapala et al. 1980). A high 

degree of host specificity has since been confirmed by others (e.g. Ahmad and Thakur 

1991, Sankaran and Sugathan 1974, Syed 1979a) and no damage to plants other than to C. 

odorata has been reported either in the Americas or in the overseas countries where it has 

become established. 

In Trinidad the eggs are parasitised by a scelionid wasp and the larvae by five 

species of tachinid fly. A nuclear polyhedrosis virus also affects the larvae (Bennett and 

Cruttwell 1973) and may have been responsible for breeding difficulties in some overseas 

countries, although other countries have experienced no problems in establishing cultures. 

Pentispa explanata Coleoptera: Chrysomelidae 

This hispine beetle is recorded on C. odoratum from Mexico to Colombia and from 

Venezuela on Pithecoctenium sp. (Bignoniaceae). In Trinidad adults are widespread on C. 

odorata and C. ivaefolia, but would not feed on Pithecoctenium echinatum. 

Eggs are inserted singly under the leaf epidermis and covered with a faecal plug. 

Larvae hatch after about 12 days and form irregular blotch mines which expand to 2 to 3 

cm in diameter 20 to 25 days later when larvae are mature. Pupation occurs in the mine 

and adults emerge 5 to 8 days later. Adults disperse and feed on the underside of the 

leaves producing characteristic scars. There is one generation a year. Larvae are para- 

sitised by a solitary ectoparasitic elasmid Austelasmus sp. and are taken by predatory 

wasps (McFadyen 1988a). 

Perasphondylia reticulata Diptera: Cecidomyiidae 

This gall fly is recorded from C. odorata and C. ivaefolia in Trinidad, Brazil and Bolivia 

and from C. odorata and Eupatorium sp. in El Salvador. 

Larvae occur singly in a hollow pear-shaped gall, 7 to 9 mm long, in stem tips and 

axillary buds. P. reticulata causes considerable damage but is generally uncommon and 

confined to the cooler valleys in Trinidad. However it is commoner in Brazil and Bolivia. 

It is attacked by several parasitoids in Trinidad and Bolivia (McFadyen 1988a). 

Procecidochares connexa Diptera: Tephritidae 

This gall fly is recorded from Mexico, Brazil and Bolivia. 

Eggs are inserted in the tip of the stem and abnormal growth commences even 

before they hatch. One to seven larvae feed in separate curved tunnels in the developing 

gall. Mature larvae pupate in the tunnel behind an epidermal window through which the 

adult emerges later. The galls slow and distort growth and cause moderate damage. 

Larvae are parasitised by a number of wasps throughout their range (McFadyen 

1988a). 

Rhodobaenus cariniventtis Coleoptera, Curculionidae 

This weevil is recorded from Trinidad and USA. Eggs are deposited in a slit between two 

rows 'of punctures encircling the stem, which result in wilting of the stem tip. On hatch- 

ing, larvae feed for a few days in the wilted portion then tunnel into the sound tissue 

below the punctures. Bennett (1955) reports that, after a month, they pupate in the stem 

at the base of the plant and adults emerge 10 days later. However, McFadyen (1988a)


states that, when mature, the larva cuts off from the hollow tip of the stem a piece about 

2cm long containing it. This falls to the ground, where the open ends are plugged with 

frass, and pupation occurs. Adults feed on stems and petioles of Bidens pilosa, 

Chromolaena inulaefolium, C. ivaefolia and C. odorata (all Asteraceae). Larvae tunnel in 

the stems of all these except B. pilosa (McFadyen 1988a). Cruttwell (1974) suggested 

that the feeding of adults might be insufficiently restricted, but the situation merits fur- 

ther investigation. 

R. cariniventris is parasitised in Trinidad by an external egg parasitoid Euderus sp. 

(Eulophidae) (Bennett 1 955). 

Comments 

The genus Chromolaena belongs to the tribe Eupatorieae (Table 4.3.4), which is mainly 

of American origin. There are no crop plants or important ornamentals in this tribe. 

However, it does contain the major weeds Ageratum conyzoides (4.1) and Mikania 

micrantha (4.14) and several less important species: Ageratina altissima in eastern USA, 

A. adenophora and A. riparia in India to southern China, Australia, Hawaii and South 

Africa and Austroeupatorium inulaefolium in Indomalaysia and Sri Lanka. There are 129 

species of Chromolaena, all from Central and South America. Chromolaena ivaefolia 

and C. laevigata are widespread and occasionally weedy in the Americas, but only C. 

odorata has spread elsewhere (McFadyen 1991a). These relationships suggest that many 

of the insects that attack C. odorata and its close allies are likely to be sufficiently host 

specific to be considered for classical biological control. 

There has been discussion concerning the possible reasons for Pareuchaetes 

pseudoinsulata establishing fairly readily in Sri Lanka and Guam, with difficulty in India 

and Sabah and not at all in Africa (e.g. Cock 1984a, Cock and Holloway 1982, Seibert 

1989). The desirability is rightly emphasised of matching, where possible, the climate of 

the area from which it (or any other biological control agent) is collected with that of the 

area in which it is to be released. Since P. pseudoinsulata has no diapause and breeds all 

year round, it will at least experience great difficulty in bridging (or find it impossible to 

bridge) the gap created by almost complete leaf loss of C. odorata in areas where there is 

a severe and long dry season. However, if this were the only problem, the moth should be 

able to establish itself at least briefly before being eliminated by starvation: this sequence 

has not, however, been documented. What, however, has been widely reported is the very 

high level of predation on eggs, larvae and pupae, in particular by ants and spiders, but 

also by other invertebrate and vertebrate predators (e.g. Kluge 1991, Kluge and Caldwell 

1991). It seems probable that massive predation has been the cause of rapid demise of 

many releases. Thus, release sites should be chosen (or treated) so as to minimise preda- 

tion. Although significant predation was also observed on Guam, the lower diversity of 

predators (and other organisms) on islands may well have contributed to the compara- 

tively ready establishment of P. pseudoinsulata there and on other Pacific islands. 

Furthermore, the release of significant numbers (500 or more) of adults rather than of 

eggs or larvae may have assisted in avoiding rapid elimination of the released material.


The general vigour of the released insects and the presence or absence of microsporida or 

viruses would also play a crucial role in successful establishment and it is probable that 

these factors have not always been adequately considered. 

It is possible that the high level of predation and of parasitisation of the biological 

control agents will, in many areas, so lessen the potential each has to cause damage to C. 

odorata that the combined effects of several will be required to bring about an adequate 

and sustained reduction in its weediness. Fortunately, many potentially suitable insects 

are available for study.




Map 4.8.1 Eleusine indica 

4.8 Eleusine indica 85 


Eleusine indica is of African origin and, except for finger millet, E. coracana, is not 

closely related to graminaceous crop plants. Finger millet is a staple crop in India and 

some parts of Africa, but relatively unimportant or not grown elsewhere. Little is known 

about the insect or other enemies of E. indica in Africa and, elsewhere, almost all records 

are of pests with a wide host range. Because it is a major weed (5th) in Southeast Asia 

and is only distantly related to crop plants a search for specific natural enemies in Africa 

must be regarded as an attractive proposition.


4.8 Eleusine indica (L.) Gaertn. 

Poaceae 

crowsfoot grass, goose grass; sin ngo let kya, sin ngo myet (Myanmar), yah 

teenka (Thailand), smao choeung tukke (Cambodia) co miin triiu (Vietnam), 

rumput sambou (Malaysia), rumput belul8ng (Indonesia), sabung sabungan 

(Philippines) 

Rating 

+++ Viet, Msia, Sing, Indo, Phil 

24 ++ Myan, Thai, Laos, Camb 

+ Brun 

Origin 

Africa (Phillips 1972), replacing an alternative view that it was India (Holm et al. 1977, 

Waterhouse 1993a). 

Distribution 

Throughout the tropics, sub-tropics and temperate regions of the world, including Africa, 

Asia, SE Asia, Australia, the Pacific and the Americas. 


E. indica is a tufted, annual, C4 grass attaining a height of 0.6 m. Its flower spikes mostly 

have 2 to 7 spikelets, producing a characteristic windmill-like appearance. 

Importance 

The genus Eleusine, contains nine annual or perennial grasses all native to Africa except 

for the South American E. tristachya (Hilu and Johnson 1992, Phillips 1972). It belongs 

to the subfamily Chloridoideae, which is but distantly related to all except one of the 

principal grain crops. That exception is finger millet, or ragi, E. coracana (2n = 36), 

which is believed to have arisen from E. indica (2n = 18) (Hilu and de Wet 1976, Hilu 

and Johnson 1992, Hiremath and Salimath 1992) and is an important staple cereal in 

India and some regions of eastern Africa (Rachie and Peters 1977). However, it is worth 

noting that E. coracana is regarded as a minor weed in some Southeast Asian countries 

(Thailand, Vietnam) (Waterhouse 1993a) and that it is nowhere important in this region. 

The genera Eleusine and Dactyloctenium are closely related. 

E. indica is an important weed in more than 60 countries in at least 46 crops and, in 

these, has the status of a serious weed in 30 countries and 27 crops. It was evaluated as 

the fifth worst weed in the world (Holm et al. 1977) and also rated fifth in a recent survey 

in Southeast Asia (Waterhouse 1993a). It was rated 15th in 1992 in the oceanic Pacific 

(Waterhouse, unpublished). It grows well in sunny or somewhat shaded places, in marsh- 

lands,. wastelands, roadsides, along borders of irrigated fields and canals, in lawns and in 

pastures, and prospers and is particularly troublesome on arable land. It ranges from near


the seashore to an altitude of at least 2000 m and is a major problem in almost all forms 

of agriculture between the tropics of Capricorn and Cancer. 

E. indica grows and flowers well in all seasons and a single plant may produce more 

than 50000 small seeds, which move readily by wind, in mud on the feet of animals and in 

the tread of machinery. The seeds are eaten by wild and domestic animals and are occasion- 

ally grown for grain in Africa and India, but Eleusine coracana, finger millet, with some- 

what larger seeds is far better for this purpose. Although sometimes claimed to be palatable 

to grazing animals, crowsfoot grass becomes fibrous too early in the season to be a satis- 

factory pasture grass. The seed heads may contain high levels of cyanogenic glycosides 

and are believed to be responsible for occasional cases of stock poisoning (Everist 1974). 


Natural enemies restricted to the genus Eleusine and its close relatives might well be con- 

sidered for biological control of E. indica except in India or other regions where finger 

millet is an important cereal. 

E. indica is reported in the literature to be attacked by more than 50 insects, nema- 

todes, fungi, bacteria and viruses, all except 6 in continents other than Africa (Tables 

4.8.1 to 4.8.4). Further, with few exceptions, all of these organisms are known to have 

wide host ranges and to attack important agricultural crops. Indeed, of those recorded, 

only one cecidomyiid gall fly and possibly one or two fungi could be considered further 

for classical biological control. Figliola et al. (1988) consider that, where they already 

occur, two fungi, Bipolaris setariae and Magnaporthe (=Pyricularia) grisea hold 

promise as bioherbicides for E. indica, although their host range is a little too wide for 

classical biological control. 

It is not surprising that the organisms attacking an economic crop, finger millet, 

E. coracana, have been investigated in greater detail than those of a weedy relative. 

Finger millet is believed to have been domesticated in the East African highlands by 

3000 BC or earlier and archaeological data suggests that it may have been introduced to 

India as early as 2000 BC (Hilu et al. 1979). Since E. coracana and E. indica are closely 

related, Wapshere (1 990b) argues, probably correctly, that most or all of the more specific 

organisms infesting finger millet are also likely to attack E. indica. It is very disappoint- 

ing, therefore, that almost all of the natural enemies of finger millet so far recorded (again 

mostly from outside Africa) have wide to very wide host ranges and are not potential bio- 

logical control agents. The very few species that may prove to have a limited host range 

are shown in table 4.8.5. Wapshere (1990b) has listed 40 insects that attack E. coracana 

and at the same time belong to groups known to have species restricted to a single grass 

genus (and there are also many other insects from groups with a wider host range that 

attack E. coracana). In addition to the undescribed Orseolia gall midge attacking E. indi- 

ca in India, only three insects (two cecidomyiid gall midges, one from Uganda and one 

from Nigeria and an aphid from India), a nematode (Heterodera delvii) from India and a 

smut fungus (Melanopsichiurn (= Ustilago) eleusinis) may, if shown to attack E. indica 

also, prove to be sufficiently host specific to be considered for classical biological control. 

It is relevant that cecidomyiid gall flies are believed to have a high degree of specificity to 

their host grass genera (Barnes 1946, K.M. Harris pers. comm. 1993, Wapshere 1990a).


Comment 

It has been pointed out above that the majority of records for natural enemies of both 

E. indica and E. coracana come from outside Africa and that almost all of these organ- 

isms have a wide host range. Indeed, this is to be expected if both Eleusine species are of 

African origin. Except for any specific enemies that may have accompanied them, it is 

inevitable that they will be attacked in new countries by non-specific natural enemies 

that, hitherto, were attacking other plants. Of course, the possibility exists that natural 

enemy species in the new country may have evolved a degree of specificity in the four or 

five thousand years that the Eleusine species have existed outside Africa. 

It is significant that there has not so far been any detailed search in Africa for natur- 

al enemies of E. indica to establish whether or not adequately specific species occur 

there. A two year (or longer) survey of E. indica in several regions of Africa would prob- 

ably be required, together with observations on whether the organisms found attacking 

E. indica also attack E. coracana, nearby grasses or other plants. The relevant regions for 

study in Africa and Madagascar are indicated in map 4.8.2 based on the distribution data 

of Phillips (1 972). 

If (i) the African cecidomyiid gall midges (Contarinia (= Stenodiplosis) spp.) 

(Tables 4.8.1, 4.8.5) do not already occur in Southeast Asia (they are not known in 

Australia), (ii) they prove to be adequately host specific and (iii) the Ugandan species 

attacks E. indica in addition to E. coracana, they would appear to be the most promising 

of known species for introduction elsewhere. The undescribed species from northern 

Nigeria (Table 4.8.1) was collected from E. indica at Zaria in July 1959 and July 1960 

(K.M. Harris, pers. comm. 1993). Larvae of the Indian Orseolia sp. nr. fluviatilis proba- 

bly induce galls on young shoots of E. indica, so would affect vegetative growth rather 

than having a direct impact on seed production. It is as yet known only from India. 

To sum up, for an attempt at classical biological control of a grass weed, E. indica 

would appear to be the one with most positive factors combined except that, so far, few 

adequately specific, natural enemies are known. However, almost nothing is known about 

the natural enemies in Africa, not only its centre of origin but also that of the genus 

Eleusine. It would, indeed, be most surprising if several natural enemies having a restrict- 

ed host range were not discovered during a thorough survey there. 

Table 4.8.1 Natural enemies of Eleusine indica: insects. 

Species Country Portion Comments: References 

attacked other hosts 

Hemiptera 

APHlDlDAE 

Chaetogeoica India 

graminiphaga 

Geoica lucijiuga India 

beans and a Raychaudhuri 1983 

number of 

grasses 

also on rice, Eleusine Raychaudhuri et al. 

coracana and many 1978 

weeds including 

Cynodon dactylon, 

Cyperus rotundus 

I (continued on next page)




Hysteroneura setariae Hawaii rice, maize, wheat 

Schizaphis (= Toxoptera) 

graminum 

Sitobion avenae 

(= Macrosiphum 

granarium) 

Sitobion (= Macrosiphum) 

miscanthi 

Sregophylla (= Anoecia) 

querci 

Tetraneura basui 

Tetraneura 

nigriabdominalis 

(= T. hirsuta) 

CERCOPIDAE 

Prosapia (= Monecphora) 

bicincta fraterna 

CICADELLIDAE 

Nephotettix malayanus 

Nephotettix 

nigromaculatus 

(= N. nigropictus) 

Nephotettix virescens 

Recilia dorsalis 

DELPHACIDAE 

Laodelphax striatellus 

sorghum, sugar cane 

rice, sorghum, maize 

and a very wide 

host range 

rice, wheat, a very 

wide host range 

India on a very wide range 

of crop plants and 

weeds 

maize and several 

weeds 

India 

on rice, Echinochloa 

colona, Paspalum 

conjugatum and 

other weeds 

India 

rice, maize, sugarcane 

Eleusine coracana 

and a very wide 

range of weeds 

Cuba also on Paspalum 

notatum, Brachiaria 

subquadripara, 

Andropogon annulatus, 


Philippines 

Philippines 

Philippines 

Philippines 





Beardsley 1962 

Patch 1939 

Patch 1939 


Patch 1939 


Patch 1939, 


Plana et al. 1986 





China wheat, barley, oats, 

sorghum etc 

Zhang et al. 1981 

Peregrinus maidis India transmitter of 

Cherian and Kylasam 

Eleusine mosaic 1937, Patch 1939, Rao 

virus (see table 4.8.4); 

very wide host range 

et al. 1965 

Sogatella furcijera China can complete 

development also on 

17 other species of 

crops and weeds 

including rice, barley, 

wheat, Echinochloa 

crus-galli 

Huang et al. 1985 






LYGAEIDAE 

Blissus leucopterus 

Thysanoptera 


Haplothrips ganglbaueri 

Diptera 

AGROMYZIDAE 

Liriomyza marginalis 

Pseudonapomyza spicata 

CEClDOMYllDAE 

Orseolia sp. nrjluviatilis 

Stenodiplosis sp. 

Lepidoptera 

ARCTllDAE 

Cnaphalocrocis medinalis 

Cnaphalocrocis 

(= Marasmia) patnalis 

Creatonotos (= Amsacta) 

gangis 

NOCTUI DAE 

Spodoptera ffugiperda 

PYRALI DAE 

Ostrinia furnacalis 

USA lower damages sorghum and Ahmad et al. 1984, 

stem many grasses including Lynch et al. 1987 

Cynodon dactylon and 

Dactyloctenium 

aegyptium, but 

particularly damaging 

to E. indica 

India rice, wheat, sorghum Ananthakrishnan & 

Thangavelu 1976 

N&S Panicum miliaceum, 

America Digitaria, Paspalum 

(primary host), 

Euchlaena 

Australia leaf maize, sugarcane, 

grasses 

India stem undescribed gall midge 

resembling (but not) 

the rice stem gall midge 

Orseolia 

(= Pachydiplosis) 

oryzae; no host other 

than E. indica known 

Nigeria seed undescribed species 

heads 

Spencer 1990, 

Spencer & Steyskal 

1986 

Kleinschmidt 1970 

Barnes 1954a,b, 1956, 

Gagnt 1985, 

Hegdekatti 1927, 

Rachie and Peters, 

1977 

K.M. Harris pers. 

comm. 1993 

Philippines leaf rice, many weeds Abenes & Khan 1990 

folder 

Philippines leaf rice, many weeds Abenes & Khan 1990 

folder 

Philippines leaves rice, many weeds Catindig et al. 1993 

USA 

Guam 

wide range of crops Pencoe and Martin 

and weeds 1982 

wide range of crops Schreiner et al. 1990 

and weeds

Table 4.8.2 Natural enemies of Eleusine indica: nematodes. 

Species 

Ditylenchus destructor 

Hirschrnaniella spinicaudata 



Meloidogyne grarninicola 



Pratylenchus pratensis 

Pratylenchus zeae 

Rotylenchulus reniformis 


Country Comments References 

South Africa 

Cuba 

China 

Cuba, 

Philippines, 

USA 

India 

Cuba, USA 

Brazil 

Hawaii 

S. Africa, 

Cuba 

Hawaii 

groundnut, several weeds 

has other weed hosts 

including Cyperus iria 

rice root knot nematode 

(damage up to 50%); also 

attacks wheat, and 

Echinochloa colona 

Echinochloa crus-galli, 

Portulaca oleracea, 

tobacco 

wheat, Panicum spp, tomato, 

capsicum, etc 

Ageratum conyzoides, Croton 

lobatus, Cynodon dactylon, 

tobacco 

attacks tomato and weeds 

including Bidens pilosa, 

Euphorbia heterophylla, 

Galinsoga parviflora 

also attacks Cynodon dactylon 

E. indica is a moderately good 

host of the maize nematode; 

has other weed hosts, including 

Cyperus iria 

Table 4.8.3 Natural enemies of Eleusine indica: fungi and bacteria. 

De Waele et al. 1990 

Femandez and Ortega 

1982 

Guo et al. 1984, Holm 

et al. 1977 

Tedford and Fortnum 

1988, Valdez 1968 

Rao et al. 1970 

Acosta et al. 1986 

Lordello et al. 1988 


Femandez and Ortega 

1982, Jordaan et al. 

1988 

Linford and Yap 1940 

Species Country Comments References 

FUNGI 

Bipolaris setariae 

(as Drechslera setariae) 

Corticium sasakii 

Drechslera gigantea 

Helminthosporium sp. 

Helminthosporium holrnii 

Helrninthosporiurn maydis 

e 

USA (not 

recorded in 

Australia) 

India 

Brazil 

Thailand 

India 

China 

heavy attack on E. indica, 

light on maize, sorghum, 

none on dicotyledons 

rice, many weeds including 

Commelina benghalensis, 


Firnbristylis miliacea 

no hosts other than E. indica 

mentioned 

also on Echinochloa colona, 

Chloris gayana 

attacks 2 1 other weeds 

including Irnperata cylindrica, 

Digitaria ciliaris and 


Figliola et al. 1988 

Hiremath and 

Sulladmath 1985 

Roy 1973 

Muchovej 1987 

Chandrasrikul 1962 

Singh and Misra 1978 

Wu and Liang 1984 




Species Country Comments References 

Helminthosporium nodulosum 

(as Bipolaris nodulosa or 

Cochliobolus nodulosus) 

Magnaporthe (= Pyricularia) 

grisea 

Pellicularia rolfsii 

Phyllachora eleusines 

Pyricularia oryzae 

Sclerophthora macrospora 

Sclerotiurn rolfsii 

Ustilago sp. 

Ustilago eleusinis (as 

Melanopsichiurn eleusinis) 

BACTERIA 

Pseudornonas glumae 

Pseudornonas plantarii 

Africa, 

Australia, 

India, Japan, 

Philippines, 

USA 

Africa, 

Australia, 

India, USA, 

Georgia 

Australia, 

India 

Africa, 

Australia 

Brazil, China 

India 

Australia, 

India 

China 

Africa, Asia 

Japan 

Japan 

also infests maize, Eleusine 

coracana, wheat, barley, oats 

and weeds including 

Dactyloctenium aegyptium; 

causes seedling blight leaf 

stripe and sooty heads in 

E. indica 

heavy attack on E. coracana, 

Rottboellia cochinchinensis, 

light attack on maize 

causes wilt disease of E. 

coracana and infests many 

grasses and dicotyledonous 

plants 

only recorded on Eleusine and 

Eragrostis in Africa; in 

Australia only on Eragrostis 

attacks rice 

attacks maize, wheat, oats, 

rice: attacks E. coracana and 

many grasses, but not E. indica 

in Australia; there may be host 

specific strains 

attacks many dicotyledonous 

crop plants and a wide range 

of grasses 

smut fungus of Eleusine and 

Dactyloctenium, but only on 

D. radulans in Australia 

an important rice pathogen: 

attacks a wide range of weeds 

attacks rice, wheat, sorghum, 

maize and many weeds 

Rachie and Peters 

1977, Wapshere 

1990b 

Chauhan & Verma 

198 1, Figliola et al. 

1988, Heath et al. 

1990, 1992, Shetty et 

al. 1985, Valent et al. 

1986, Vodianaia et al. 

1986, Wapshere 

1990b,c 

Wapshere 1990b 

Parbery 1967, 

Ramakrishran 1963 

Prabhu et al. 1992, 

Teng 1932, Valent et 

al. 1986 

Rachie and Peters 

1977, Ullstrup 1955, 

Wapshere 1990b 

Reddy 1983, Safeeulla 

1976 

Mundkur 1939 

Simmonds 1966, 

Zundel 1953 

Miyagawa et al. 1988 

Miyagawa et al. 1988

Table 4.8.4 Natural enemies of Ebusine indica: viruses. 

Virus Country Other hosts 

cereal chlorotic mottle 

corn leaf gall 

corn stunt 

Eleusine mosaic 


maize dwarf mosaic 

maize streak 

rice leaf gall 

rice orange leaf 

rice ragged stunt 

rice tungro bacilliform 

rice tungro spherical 

rice yellow mottle 

sugarcane mosaic 

sugarcane streak 

tungro 

wheat rosette 

Australia 

Philippines 

USA 

India 

Malawi 

USA 

Nigeria 

Philippines 

Philippines 

China 

Philippines 

Philippines 

Philippines 

Kenya 

India 

Hawaii 

Philippines 

China 

oats, barley, wheat, maize, 

E. coracana, Digitaria ciliaris, 

Echinochloa colona; 

transmitted by Nesoclutha 

pallida 

maize 

several other weeds 

maize, sorghum, E. coracana 

and many other hosts 

groundnut 

maize 

maize, but not all cultivars 

Cicadulina triangula is 

the vector 

rice 

rice 

rice, E. indica and 4 other 

weeds 

rice, Echinochloa 

glabrescens, Monochoria 

vaginalis, Paspalum distichum 



rice, two grasses 

sugarcane 

sugarcane 

rice 

oats, barley, sorghum, 

wheat etc. Laodelphax 

striatellus is the vector 


References 

Greber 1979 

Agati and Calica 1950 

Pitre and Boyd 1970 

Rao et al. 1965 

Adams 1967 

Lee 1964 

Ekukole et al. 1989, 

Rossel et al. 1984 

Agati and Calica 1950 

Watanakul 1964 

Xie et al. 1984 

Salamat et al. 1987 



Okioma et al. 1983 

Chona and Rafay 1950 


Watanakul 1964 

Zhang et al. 1981 

Table 4.8.5 Natural enemies of Eleusine coracana which may prove to have 

a limited host range. 

Species Country Portion Comments References 

attacked 

INSECTS 

Diptera 

CEClDOMYllDAE 

Contarinia sp. Uganda inflores- not the same as the 

cence sorghum midge Barnes 1946, 1954a,b, 

Contarinia sorghicola: 1956, Geering 1953, 

the same or a similar Rachie & Peters 1977 

species attacks the 

common fallow weed 

Sorghum verticilliflorum 




Species Country Portion Comments References 

attacked 

Hemiptera 

APHlDlDAE 

Sitobion 

(= Macrosiphum) 

leelarnaniae 

NEMATODES 

Heterodera delvii 

FUNGI 

Melanopsichium 

eleusinis(= Ustilago 

eleusinis) 

India attacks several millets Raychaudhuri 1983 

(not in in India including pearl 

Australia) millet Pennisetum 

glaucum (=P. typhoideum), 

also Andropogon vulgare 

India root no other hosts Jairajpuri et al. 1979 

cysts mentioned 

Asia, Africa a smut fungus: only Simmonds, 1966, 

from Eleusine and Wapshere 1990c, 

Dactylocteniurn: Zundel 1953 

tentatively identified 

from D. radulans in 

Queensland, but not 

from E. indica

4.8 Eleusine indica 

Map 4.8.2 Distribution of Eleusine indica in Africa (after Phillips 1972)



(after Barnes and Chan, 1990)

Map 4.9 Euphorbia heterophylla 

4.9 Euphorbia heterophylla 97 


There are very few records of natural enemies other than fungi attacking Euphorbia 

heterophylla and no study has been made in tropical America where it evolved. However, 

from the sparse records of insects attacking species of Euphorbia in Brazil it is likely that 

adequately host specific insects do occur. Nevertheless E. heterophylla is regarded as an 

important weed in southern Brazil.


4.9 E uphorbia heterophylla L. 

(= E. geniculata = E. prunifolia) 

Euphorbiaceae 

painted spurge, Mexican fire plant; yaa yaang (Thailand) 

Rating 

+++ Thai 

10 ++ Msia 

+ Myan, Laos, Camb, Viet, Phil 

Indo 

Origin 

Tropical and sub-tropical America. 

Distribution 

Widespread as a weed in the tropical and subtropical regions of the world, notably in 

Southeast Asia, but apparently not in Kalimantan or Sulawesi (Indonesia) (Soerjani et al. 

1986). 


An erect annual, up to about 1 m tall; stem cylindrical, hairy; lower leaves alternate; 

stems and leaves with milky latex. The simple or lobed leaves are crowded towards the 

top of the stem, with a flat, dichotomously-branched, terminal inflorescence of small yel- 

low flowers and large leafy bracts, often with a bright red or cream patch at the base. The 

inflorescence consists of many small, short-stalked flowers lacking petals but with con- 

spicuous glands (Wilson 1981). Reproduction is by seeds which are shed with an explo- 

sive mechanism. 

Importance 

A weed of increasing importance in upland fields of rice and many other crops; also in 

wastelands, roadsides, boundaries of coffee plantations; very abundant locally. Seeds per- 

sist in the soil until favourable conditions allow germination and rapid growth, giving 

rise to large populations of the weed. It is an important weed in 23 tropical countries and 

present in at least 37 others. Its rapid growth enables it to compete successfully with 

crops, quickly forming a dense canopy over young crop plants. Dense populations of the 

weed, with its white sticky latex, may make it impossible to harvest the crop. 

The young leaves are sometimes used as a vegetable, but are laxative if too much is 

eaten. The plant is said to have caused poisoning in livestock (Wilson 1981). 


Except possibly for Alternaria sp. and Helminthosporium sp. which have not been shown 

to be pathogenic to crop plants (Yorinori 1985), there are no records of apparently host


specific organisms attacking Euphorbia heterophylla (Table 4.9.1). However, it is known 

that a number of insects do attack it in Brazil, but this observation was incidental to a 

study of fungi and none of the insects were identified (E.G. Fontes, pers. comm. 1992). 

Although periodic collections were made in Trinidad in the early 1970's, no promising 

insects were encountered (Yaseen 1972). 

There are few records (19 only) of insects attacking members of the genus 

Euphorbia in Brazil (Table 4.9.2) (d'Araujo e Silva et al. 1968a,b), indicating that little 

attention has so far been paid to Euphorbia spp. in this region. Six of the insects are 

polyphagous and too little is known about the others to arrive at a conclusion. Even if 

some are restricted to the Euphorbiaceae, it remains to be determined whether any will 

attack either Euphorbia heterophylla or E. hirta. 

E. heterophylla is resistant to most herbicides and, in recent years, has become pro- 

gressively more important in Brazil, particularly in the southern, soybean-producing 

states (Yorinori 1985), which suggests that its insect enemies, if any, may be heavily par- 

asitised. 

A biological control program has been in progress in Canada since the late 1960's 

against Euphorbia cyparissias and E. pseudovirgata, involving the introduction of some 

twenty species of insects from Europe. Several species have become established, with 

rather localised effects (Julien 1992). It is said that insects are generally unable to attack 

Euphorbia species because of the latex that flows freely from any wound and clogs the 

mouthparts (Best et al. 1980), but clearly some insects are adapted to deal with this prob- 

lem. 

The best known economic plant in the Euphorbiaceae is cassava, Manihot esculenta 

of South American origin. The insects attacking it there are comparatively well known, a 

factor that will aid the evaluation of the specificity of insects attacking Euphorbia spp. 

Another species of horticultural importance is poinsettia, Euphorbia pulcherrima. 

Table 4.9.1 Natural enemies of Euphorbia heterophylla. 


INSECTS 

Orthoptera 

ACRlDlDAE 

Poekilocerus 

hieroglyphicus 

Hemiptera 

ALEYRODIDAE 


ALYDIDAE 

Leptocorisa acuta 

Leptocorisa oratorius 

Leptocorisa 

solomonensis 

Sudan beans, melons Ba-Angood 1977, 

Ba-Angood & 

Khidir 1975 

Thailand, cotton, polyphagous Debrot & Centeno 1985, 

Venezuela Nachapong & Mabbett 1979 

PNG 

PNG 

PNG 

F. Dori pers. comm. 1993 






MITES 

FUNGI 


TETRANYCHIDAE 

Tetranychus urticae Cuba polyphagous Perez et al. 1987 

Alternaria sp. 

Amphobotrys ricini 

Elsinoe sp. 

Helminthosporium sp. 

Macrophornina 

phaseolinu 

Phytophthora 

palmivora 

Puccinia sp. 

Rhizoctonia solani 

Sclerotinia 

sclerotiorum 

Sphaceloma sp. 

Uromyces euphorbiae 

NEMATODES 

Meloidogyne exigua 


Rotylenchulus 

reniformis 

Brazil 

USA 

Burundi 

Brazil 

India 

cassava 

Sarawak black pepper Anon 1979 

Brazil 

Brazil 

Brazil 

Brazil, 

Burundi 

Brazil 

Brazil 

Brazil 

Florida 

VIRUSES 

Euphorbia mosaic Brazil, USA, 

Venezuela 

cassava 

Yorinori 1985 

Holcomb et al. 1989 

Zeigler & Lozano 1983 

Fontes et al. 1992, 

Gazziero et al. 1988, 

Yorinori 1985 

Saxena et al. 198 1 

Fontes et al. 1992 

Yorinori 1985 

Yorinori 1985 

Yorinori 1985, 

Zeigler & Lozano 1983 

Yorinori 1985 

coffee, many weeds Luc et al. 1990 

Lordello et al. 1988 

Inserra et al. 1989, 

MacGowan 1989 

Debrot & Centeno 1985, 

Kim & Flores 1979, 

Kim & Fulton 1984, 

Yorinori 1985 

Table 4.9.2 Insects attacking species of Euphorbia in Brazil (d'Araujo e Silva et al. 

1968a,b). 

Insect Hosts Feeding habit 

Hemiptera 

ALEYRODIDAE 


(= B. costa-limai) Euphorbia hirtella, polyphagous 

tomato, Mentha arvensis 

COCCIDAE 

Coccus spp. Euphorbiaceae, Acalypha sp., polyphagous 

Aspidosperma ramiflorum, 

Cassia sp., Citrus spp. 



Insect Hosts Feeding habit 

Eucalymnatus spp. 

Platinglisia noacki 

COREIDAE 

Chariesterus armatus 

TlNGlDAE 

Corythuca pellucida 

Corythuca socia 

Thysanoptera 


Haplothrips gowdeyi 

Coleoptera 

CHRYSOMELIDAE 

Caryedes (= Gibbobruchus) 

pickeli 

Disonycha argentiniensis 

Gibbobruchus polycoccus 

CURCULIONIDAE 

Sternocoelus sp. 

Sternocoelus notaticeps 

Lepidoptera 

LYMANTRIIDAE 

Thagona tibialis 

NOCTUIDAE 


NYMPHALIDAE 

Didonis biblis 

Dynamine artemisia 

Episcada pascua 

SPHlNGlDAE 

Erinnyis oenotrus 

Euphorbia capansa, Nerium sp., polyphagous 

Caryota sp., Phoenix sp. 

Euphorbiaceae, Begonia sp., polyphagous 

Eugenia sp., Grevillea robusta, 

Ilex sp., kurus sp., Magnolia 

pumila, etc. 

Euphorbia braziliensis 

Euphorbiaceae 

Euphorbiaceae 

Euphorbia sp., coffee, rice, 

Crotolaria sp., PassiJlora sp., 

Buddleia variabilis 

Euphorbiaceae 

Euphorbia pulcherrima 

Euphorbiaceae 

Euphorbiaceae 

Euphorbiaceae 

E. cespitosa, E. ovalifolia, 

E. pulcherrima 

Euphorbiaceae, many crops 

Euphorbiaceae, Tragia volubilis 

Euphorbiaceae 

Euphorbiaceae 

E. ovalifolia 

possibly restricted 



polyphagous 




?restricted to Euphorbiaceae 



polyphagous 




possibly restricted



(after Holm st a/. 1977)

Map 4.10 Euphorbia hirta 

4.10 Euphorbia hirta 


There is only one record of a natural enemy attacking Euphorbia hirta in tropical 

America where it evolved and only a few of polyphagous species attacking it elsewhere. 

A survey in Central America would be necessary to determine what species attack it 

there that might be potential biological control agents.

104 


4.10 Euphorbia hirta L. 

(= E. pilulifera) 

Euphorbiaceae 

garden spurge, asthma plant; mayo (Myanmar), nam nom raatchasee (Thailand) 

tuk das khla thom (Cambodia), co sua 16ng (Vietnam), ara tanah, hairy spurge 

(Malaysia) gelang susu, gendong ancok (Indonesia), gatas gatas (Philippines) 

Rating 

++ Thai, Sing, Phil 

10 + Laos, Camb, Viet, Msia 

w Myan, Indo 

Origin 


Distribution 

E. hirta is a weed of the tropics and subtropics. 


A small, prostrate, hairy annual, 0.15 to 0.3 m tall, with a tap root; stems much branched 

from the base, often reddish, bearing brownish stiff hairs and having milky sap; leaves, 

hairy, opposite, sometimes purple-blotched and with toothed margins; flowers unisexual; 

reproduction by seeds 0.5 to 1 mm long. 

Importance 

E. hirta grows well in sunny to lightly shaded cultivated lands, gardens, lawns, waste 

areas and run down grasslands. It is an early coloniser of bare ground especially under 

damp or irrigated conditions. It flowers all year round in Southeast Asia producing up to 

3000 seeds per plant. When the seed pods mature they explode to disperse the seeds. Its 

prostrate habit enables it to tolerate mowing and it can be important in lawns. It has been 

reported from 47 countries as a weed in many crops, including citrus, cotton, groundnuts, 

maize, pineapples, rice, sorghum, sugarcane, tea and vegetables. Moody (1 989) records it 

as being more widespread in rice than Euphorbia heterophylla. 

E. hirta is sometimes used in medicines in Fiji, Malaysia, Indonesia, the Philippines 

and Brazil-the leaves and latex against intestinal diseases, ulcers and bronchitis, and the 

latex for conjunctivitis. It may have slightly poisonous properties and is useless as fodder 

for livestock. 


In view of its common occurrence in the tropical and subtropical belt of the world, it is 

surprising that there are so few records of natural enemies attacking it, and those that do 

are highly polyphagous (Table 4.10.1). A survey in Central America would be necessary 

to learn more about its natural enemies that might have potential for biological control.

Table 4.10.1 Natural enemies of Euphorbia hirta. 

4.10 Euphorbia hirta 105 


INSECTS 

Orthoptera 

ACRlDlDAE 

Chrotogonus 

trachypterus 

Hemiptera 

APHlDlDAE 

Aphis craccivora 


ALEYRODIDAE 


DELPHACIDAE 

Tarophagus proserpina 

LYGAEIDAE 

Nysius inconspicuus 

PSEUDOCOCCIDAE 

Ferrisia virgata 

Thysanoptera 

THRlPlDAE 

Haplothrips euphorbiae 

Diptera 

AGROMYZIDAE 

Liriomyza bryoniae 

Liriomyza strigata 

Lepidoptera 

NOCTUI DAE 

Achaea janata 

FUNGI 

Aecidium tithymali 

Amphobotrys ricini 


quinqueseptatum 

Phytophthora palmivora 

PROTOZOA 

Phytomonas sp. 

NEMATODES 


India 

Nigeria, 

Uganda 

India 

India 

Philippines 

India 

India 

India 

Europe 

W. Europe, 

USSR 

Indonesia 

Thailand 

USA 

India 

Sarawak 

Venezuela 

Hawaii 

polyphagous Chandra et al. 1983 

poly phagous, Booker 1964, Davies 1972, 

a virus transmitter Ofuya 1988 

polyphagous Jeritta & David 1986 

polyphagous, Jeyarajan et al. 1988 

a virus transmitter 

polyphagous Duatin & Pedro 1986 

polyphagous Thangavelu 1978 

poly phagous Jeritta & David 1986 

possibly host restricted Jeritta & David 1986 

highly polyphagous Spencer 1973, 1990 

highly polyphagous Spencer 1973, 1990 

polyphagous Kalshoven 198 1 

Puckdeedindan 1966 

Holcomb et al. 1989 

Sulochana et al. 1982 

black pepper Anon 1979 

Barreto 1982 

Valdez 1968 





Meloidogyne javanica India Dahiya et al. 1988 

Radopholus sirnilis Zimbabwe polyphagous Martin et al. 1969 

Rotylenchulus reniforrnis Hawaii, USA Linford & Yap 1940, 

Inserra et al. 1989 

VIRUSES 

groundnut rosette Hawaii, Nigeria, Adams 1967, 

Uganda Booker 1964, 

Davies 1972 

hibiscus yellow India Jeyarajan et al. 1988 

vein mosaic 

tapioca mosaic India Jeyarajan et al. 1988 

tobacco leaf curl Hawaii Holm et al. 1977 

tomato leaf curl India Jeyarajan et al. 1988 

urd bean yellow mosaic India Jeyaragan et al. 1988




Map 4.11 Fimbristylis miliacea 

4.1 1 Fimbristylis miliacea 109 


Very few natural enemies of Fimbristylis miliacea are known and it would be necessary 

to carry out a survey in tropical America before it would be possible to evaluate the 

prospects for its biological control.


4.11 Fimbristylis miliacea (L.) Vahl 

(= l? littoralis) 

Cyperaceae 

lesser fimbristylis, grass-like fimbristylis; m6nhnyin (Myanmar), kak phrek kdam, 

smao (Cambodia), rumput bukit, rumput tahi berbau (Malaysia), agor (Thailand), 

ba bawagan (Indonesia) 

Rating 

+++ Myan, Camb, Msia, Brun, Indo 

23 ++ Viet, Sing, Phil 

. Thai, Laos 

Origin 


Distribution 

Central America, West Africa, Asia and Southeast Asia to northern Australia. 


An erect annual or perennial sedge, growing up to 0.9 m; flower stems 4 or 5 angled, 

leaves two-ranked, threadlike, stiff and half as long as flower stems; inflorescence a dif- 

fuse compound umbel. 

Importance 

F. miliacea thrives in damp, open waste places, competing actively with other vegetation 

following germination during dry periods or shallow water conditions. A layer of water 

15 cm deep suppresses germination. Seedlings emerge during the entire growing period 

of rice with which it competes actively. It is a troublesome weed in 21 countries. In 

Malaysia it is the first sedge to emerge after rice has been transplanted and the first to 

recover after ploughing. In the Philippines it flowers all year, plants each producing up to 

10000 seeds. In many places there is no seed dormancy. 

F. miliacea is one of the most serious and widespread weeds of rice and is also 

reported from taro (Hawaii), bananas (Taiwan), abaca (Philippines), maize, sugarcane 

(Indonesia, Taiwan) and sorghum (Malaysia). 

F. miliacea is eaten by cattle, but the seeds are mostly undigested and germinate 

near the dung. 

~atu'ral enemies 

So little is known about its natural enemies (Table 4.1 1.1) that it is not possible to evalu- 

ate the prospects for biological control. A survey is necessary in tropical America.

Table 4.11.1 Natural enemies of Fimbrisstylis miliacea. 

4.1 1 Fimbristylis miliacea 111 


INSECTS 

Hemiptera 

PENTATOMIDAE 

Scotinophara Philippines rice, Cornrnelina Barrion & Litsinger 1987 

latiuscula benghalensis, 


Lepidoptera 

Creatonoros gangis Philippines rice, many weeds Catindig et al. 1993 

(= Amsacra) 

FUNGI 

Corticum sasakii India Commelina Roy 1973 

benghalensis, 


Eleusine indica and 

other grasses 

NEMATODES 

Criconemella onoensis rice, many weeds Luc et al. 1990 

Hirschmaniella spp. rice, maize, sugarcane, Luc et al. 1990 

many weeds 

Meloidogyne India rice, many weeds Luc et al. 1990, 

graminicola Rao et al. 1970 

Meloidogyne oryzae Surinam rice, plantain, wheat, Maas et al. 1978 

potato, tomato 

Rotylenchulus Trinidad very polyphagous Singh 1974 

reniformis


Marsilea minu fa 

(after Soerjani el a/. 1987)

Map 4.12 Marsilea minuta 

4.1 2 Marsilea minuta 113 


Marsilea minuta, water clover, is thought to be of tropical African origin, but no account 

of its natural enemies there is available. A survey would thus be required to evaluate the 

prospects for its biological control.


4.12 Marsilea minuta L. 

(=M. crenata) 

Marsileaceae 

water clover, clover fern, pepperwort; pak vaen (Laos), chuntul phnom 

(Cambodia), semanggi (Indonesia), phak waen (Thailand), tapah itik (Malaysia) 

paang itik (Philippines). 

Rating 

+++ Msia 

12 ++ Indo, Phil 

+ Myan, Thai, Laos, Camb, Viet 

Origin 

Africa or possibly tropical Asia (Jacobsen 1983). It consists of a complex of strains 

including a diploid (n = 20) and a sterile triploid (2n = 60) (Tryon and Tryon 1982). 

Distribution 

Marsilea minuta is widespread over most of the African continent and it is pantropical in 

Asia. The Marsileaceae contains about 65 species, of which 16 occur in Africa and, of 

these, M. minuta is amongst the most widespread (Jacobsen 1983). 


A very variable, perennial water fern of aquatic or marshy sites. Its stems are creeping 

rhizomes rooted in the mud. Leaves are clover-like, with four leaflets borne on a petiole 2 

to 30 cm long. Leaflets have fan-shaped, repeatedly bifurcating veins and normally float 

on the water surface. Sporocarps occur near the base of the petioles and usually occur 

under the mud or water surface. Reproduction is by spores or rhizomes. 

Importance 

Although M. minuta has a rating of 12 and is widely reported as a weed in Southeast 

Asia, there are surprisingly few references to it as a weed in the literature except for 

those dealing with chemical control. In Thailand it is common in rice fields and along 

canals and other waterways. It is one of the seven most important weeds in the Muda 

area of Malaysia (Itoh 1991a). It is one of the more important emergent weeds in shallow 

water rice fields in the central lowlands in Vietnam (Nguyen Van Vuong 1973) and in the 

lowland area of Vientiane in Laos (Sisounthone and Sisombat 1973). In Indonesia its 

vegetative growth and reproduction is very rapid. It can grow under water and, after 

weeding, rapidly re-establishes itself unless well buried in the soil. It is an effective com- 

petitor for nutrients, particularly in the early part of the growth period after transplanta- 

tion of rice seedlings when M. minuta rapidly covers the ground surface. In the 

Philippines it caused 19% crop loss when sown together with rice (Suriapermana 1977). 

The young leaves of water clover are sometimes eaten as a vegetable in Indonesia.


4.1 2 Marsilea minuta 115 

Very little information concerning natural enemies emerged from computer-aided search- 

es of the literature on Marsilea minuta, which also included searches of its synonyms: 

M. crenata and M. erosus in Asia and M. difisa, M. perrieriana and M. senegalensis in 

Africa (Table 4.12.1). In Africa Marsilea minuta appears to be regarded, at most, as a 

minor weed. This is possibly due to the fact that rice is far less important there than in 

Asia, or it may be due to the presence of effective natural enemies in Africa, although 

these have not yet been reported. 

In the Philippines, larvae of the ephydrid flies Notiphila Eatigenis and N. similis are 

common on emergent M. minuta and damage its stems. Their eggs are usually laid on the 

stems and serve as alternative hosts of Trichogramma wasps attacking rice stem borers 

(Barrion and Litsinger 1986). The contents of upwards of 90% of the sporocarps from 

M. minuta growing under terrestrial conditions in northwestern India were destroyed by 

larvae of the weevil Echinocnemus. The larval and pupal stages are completed in 40 to 45 

days (Loyal and Kumar 1977). In Indonesia the case-forming larvae of the widely distrib- 

uted pyralid moth Elophila (= Nymphula) responsalis attacked M. minuta and several 

other aquatic plants including Salvinia spp., Lemna purpusilla, L. polyrhiza, Monochoria 

vaginalis, Azolla pinnata and Pistia stratiotes. However, tests have shown that it will not 

feed on rice. The development period of Elophila responsalis ranged from 42 to 56 days. 

It was attacked by a pupal parasitoid (Tetrastichus sp.) and a larval coleopteran predator 

(Handayani and Syed 1976, Sankaran and Rao 1972, Subagyo 1975). Elophila respon- 

salis occurs also in India, Sri Lanka, Myanmar, Japan and Australia. 

Comment 

A survey for natural enemies attacking M. minuta in Africa is required before the 

prospects for its biological control in Southeast Asia can be evaluated. 

Table 4.12.1 Natural enemies of Marsilea minuta. 

Species Location References 

INSECTS 

Coleoptera 

CURCULIONIDAE 

Echinochnemus sp. 

Diptera 

EPHYDRIDAE 

Notiphila latigenis 

Notiphila similis 

India Loyal & Kumar 1977 

Philippines Barrion & Litsinger 1986 

Philippines Barrion & Litsinger 1986 





Lepidoptera 

PY RALl DAE 

Elophila (= Nymphula) 

responsalis 

FUNGI 

VIRUS 

Alternaria sp. 

Cercospora rnarsileae 

Phaeotrichoconis crotalariae 

Pistia virus 

India, Indonesia 

India 

India 

India 

India 

Handayani & Syed 1976, 

Sankaran & Rao 1972, 

Subagyo 1975 

Menon & Ponnappa 1964 

Patil 1975 


Menon & Ponnappa 1964



(after Soerjani et a/. 1987)

Map 4.13 Melastoma malabathricum 

4.1 3 Melastoma malabathricum 119 


M. rnalabathricum is a perennial shrub which probably originated in Southeast Asia or 

neighbouring areas, including Irian Jaya, Papua New Guinea and northern Australia, a 

region where it is regarded as being of little importance. A survey in this region would 

reveal whether there are promising natural enemies for biological control.


4.13 Melastoma malabathricum L. 

(= Melastoma affine) 

Melastomataceae 

melastoma, Indian rhododendron, Straits rhododendron; senduduk (Malaysia) 

Rating 

+++ Msia, Brun 

13 ++ Sing 

+ Thai, Laos, Viet, Indo, Phil 

Camb 

Origin 

Asia, Papua New Guinea, Australia. 


M. malabathricum is a perennial shrub growing to 2 m high; its stems are reddish with 

rough upwardly pointing scales, the leaves are tapered to both ends, are rough to touch 

and have three distinct ribs. The flowers, which are clustered at the ends of twigs, are 

pinkish to light violet. The fruit is a berry-like capsule covered with scales. 

Importance 

Melastoma is common in abandoned clearings, on waste ground and in cultivated lands. 

In addition to its importance in Southeast Asia, it is a principal weed of rubber in West 

Africa. It is said to make good firewood. The sweetish black seeds are eaten and chewed 

leaves are used for burns and against amoebic dysentery. The fruits host a fruit fly 

species in the Bactrocera dorsalis complex which does not attack commercially impor- 

tant fruits (R.A.I. Drew, pers. comm.). 


Krauss (1 965) surveyed the natural enemies of species of Melastoma, including M. mala- 

bathricum, in various countries of Southeast Asia and islands of the western Pacific. 

Twenty six insect species were found on M. malabathricum (Table 4.13.1) and a further 

34 species on other melastomas. It is very likely that some of the 34 species will also 

attack M. malabathricum and, indeed, the leaf rolling pyralid moth Ategumia fatualis 

does so. After specificity tests Ategumia fatualis was liberated in Hawaii and Kauai in 

1958 and became established, although it did not become sufficiently abundant to pro- 

vide effective control (Table 4.13.2) (Krauss 1965). 

Another leaf-rolling pyralid Ategumia adipalis was liberated in 1965, and was 

reported to have become established (Davis and Chong 1969), but at low population lev- 

els. Next a noctuid moth Selca brunella was introduced to Kauai and Hawaii from 

Malaysia and Singapore in 1964 and was recovered the next year. The larvae feed avidly

4.13 Melastoma malabathricum 121 

in flower buds, bore into terminal stems and eat leaves. In heavily infested localities con- 

siderable dieback has resulted, at places flowering was prevented and in others up to 50% 

of fruits were destroyed by larvae. Larvae have been found recently on Tiboochina urvil- 

leana and Heterocentron subtriplinenium (both Melastomataceae) in Hawaii (C.J. Davis 

pers. comm. 1993). A braconid wasp Meteorus sp. attacks S. brunella larvae (Davis 

1970, Davis and Chong 1969). 

An unidentified grasshopper and an unidentified lepidopterous larva attack M. mal- 

abathricum in Thailand but not the chrysomelid beetle Altica cyanea which is present 

there and attacks it in Indonesia and Malaysia (Napompeth 1982). 

Comment 

Although a number of insects are known to attack M. malabathricum in Southeast Asia 

(and especially in Malaysia), they clearly do not reduce its status to the level required and 

thus are of limited value for classical biological control in that region. However, if as 

postulated, the area of origin includes Papua New Guinea, (Irian Jaya) and Australia it is 

possible that there may be useful natural enemies in the region that do not occur in coun- 

tries to the north and west. Certainly, Melastoma is not listed as an important weed in 

Papua New Guinea. 

Table 4.13.1 Natural enemies of Melastoma malabathricum. 

Species Country Food References 

INSECTS 

Hemiptera 

APHlDlDAE 

Aphis sp. 

CICADELLIDAE 

Tettigella 

(= Tettigoniella) sp. 

COCCI DAE 

Rastrococcus sp. 

MEMBRACIDAE 

Gargara sp. 

kptocentrus taurus 

Nilaurama minutispina 

Sipylus dilatatum 

Sipylus sp. 

Tricentrus sp. 

MlRlDAE 

Helopeltis antonii 

Hyalopeplus vitripennis 

RlCANllDAE 

- Pochazia antica 

Singapore leaf 

Malaysia leaf 

Malaysia 

Malaysia 

Malaysia 

Malaysia 

Malaysia 

Malaysia 

Malaysia 

Singapore 

Indonesia 

Malaysia 

Malaysia 

leaf 

branch; also on 

Melastoma 

polyanthum 

branch 

branch 

branch 

leaf 

leaf 

Krauss 1965 

Krauss 1965 

Krauss 1965 

Krauss 1965 

Krauss 1965 

Krauss 1965 

Krauss 1965 

Krauss 1965 

Krauss 1965 

Krauss 1965 

Soerjani et al. 1987 

Krauss 1965 

Krauss 1965 




Species Country Food References 

Coleoptera 

CHRYSOMELIDAE 

Altica cyanea Indonesia, 

CURCULIONIDAE 

Alcidodes sp. 

Ceutorhynchus sp. 

Cryptorhynchus sp. 

Imerodes (?) sp. 

Malaysia 

Malaysia 

Malaysia 

Malaysia 

Singapore 

Diptera 

TEPHRITIDAE 

Bactrocera dorsalis 

(= B. pedestris) Malaysia, 

Singapore, 

Sri Lanka 

Lepidoptera 

ARCTllDAE 

Species of Lithosiinae 

GELECHllDAE 

Idiophantis sp. 

HYPONOMEUTIDAE 

Argyresthia leuculias 

LYMANTRIIDAE 

?Species 

NOCTUIDAE 

Autoba (= Eublemma) 

versicolor 

Selca brunella 

PY RALl DAE 

Agrotera basinotata 

Ategumia adipalis 

Ategumia fatualis 

TORTRICIDAE 

Archips rnicaceana 

shoot 

flowers 

flowers 

flowers 

Malaysia fruit 

Malaysia 

Malaysia 

Malaysia 

Malaysia 

Malaysia, 

Singapore 

Malaysia 

Malaysia, 

Singapore 

Philippines 

Kamarudin & Shah 

1978, Napompeth 1982 

Krauss 1965 

Krauss 1965 

Krauss 1965 

Krauss 1965 

fruit Krauss 1965 

fruit 

fruit 

flower 

leaf 

leaf, twig, fruit 

leaf 

leaf 

Krauss 1965 

Krauss 1965 

Krauss 1965 

Krauss 1965 

Krauss 1965 

Julien 1992, Krauss 1965 

Krauss 1965 

Julien 1992, Krauss 1965 

leaf Krauss 1965 

Malaysia leaf Krauss 1965 

FUNGI 

Phytophthora palmivora Sarawak black pepper Anon 1979

4.13 Melastoma malabathricum 123 

Table 4.13.2 Introductions to Hawaii for the biological control of Melastoma 

malabathricum. 

Species Source Liberated Established References 

INSECT 

Lepidoptera 

NOLIDAE 

Selca brunella 

PYRALl DAE 

Ategumia adipalis 

Ategumia fatualis 

Malaysia, 1965 + 

Singapore 

Malaysia, 1965 + 

Singapore 

Philippines 1958 + 

Davis 1960, Davis & 

Chong 1968, Davis & 

Krauss 1962, 1966, 

1967, Julien 1992, 

Krauss 1965 

Davis & Chong 1969, 

Julien 1992 

Davis & Krauss 1966, 

Julien 1992




Map 4.14 Mikania micrantha 

4.1 4 Mikania micrantha 125 


Mikania micrantha is a perennial vine, native to Central and South America. 

A number of very promising, and probably specific, natural enemies are known in 

Central and South America where M. micrantha is not regarded as a weed. One of these, 

a thrips Liothrips mikaniae has been released in Malaysia and the Solomon Is, but 

extremely high predation is believed to have prevented its establishment. A bug 

Teleonemia sp., several chrysomelid beetles and an eriophyid mite Acalitus sp. warrant 

serious consideration. A number of other natural enemies, whose specificity has not yet 

been adequately investigated, also attack M. micrantha. 

In spite of the lack of success with the thrips, M. micrantha appears to be a prime 

target for the introduction of one or more of the other organisms that attack it in its area 

of origin.


4.14 Mikania micrantha Kunth 

Asteraceae 

mile-a-minute weed; cheroma, ulam tikus (Malaysia), sembung rambat 

(Indonesia). 

This chapter updates that in Waterhouse and Norris (1987), with special reference to 

Southeast Asia. 

Rating 

+++ Msia, Sing 

11 ++ Brun, Indo 

+ Phil 

Thai 

Origin 

The weedy species in Southeast Asia and the Pacific is M. micrantha from Central and 

South America and not the North American M. scandens or the Old World M. cordata 

(Parker 1 972). 

Distribution 

M. micrantha belongs to a genus containing about 250 species of mostly Central and 

South American origin. In addition to its native distribution in tropical America, it has 

spread to Mauritius, India, Sri Lanka, Bangladesh and Southeast Asia (as above). It 

occurs widely as a weed in the Pacific, including Papua New Guinea (Waterhouse and 

Norris 1987), but is not yet present in Australia. It was introduced from Paraguay to 

Bogor Botanic Gardens (Indonesia) in 1949 and, in 1956, was used as a soil cover in rub- 

ber: it has since spread throughout Indonesia (Soerjani et al. 1987). 


Mikania micrantha is an extremely fast growing, sprawling, perennial vine, with oppo- 

site, heart-shaped leaves, longitudinally ribbed, branched and hairless stems and numer- 

ous small heads of densely clustered white flowers. It creeps and twines, roots readily at 

the nodes and produces abundant small (2 mm long) black seeds bearing a terminal tuft 

of white bristles that aid wind dispersal. 

In its natural habitat in tropical America, M. micrantha is usually found in disturbed 

situations. It seldom occurs on poor soils and is most commonly found in damp or 

swampy places. Typical sites in South America are roadsides in wet forest and the edges 

of freshwater swamps. Flowering occurs mainly in the dry season and only in sunny situ- 

ations (Cock 1982a). 

Importance 

With its rapid growth, ready rooting at nodes, smothering habit and prolific seed produc- 

tion, M. micrantha rapidly colonises disturbed habitats, retarding, by competition and

4.14 Mikania micrantha 127 

through plant inhibitors that it elaborates (Wong 1964), the growth of crops or natural 

vegetation. In comparison with a nitrogen-fixing legume, it is of restricted value in the 

role of a cover crop. For example, in Malaysia the girth of rubber trees was 27% less 

with a cover of M. micrantha than of a legume and the yield over the first 32 months of 

production was 27 to 29% less (Teoh et al. 1985). In many parts of Southeast Asia it is a 

serious pest of plantation crops (oil palm, coconut, cocoa, tea, rubber, teak). Its climbing 

habit enables it to reach and then dominate the crowns of bushes or trees up to 10m high, 

where it is difficult to attack either mechanically or chemically without risk of damaging 

the crop (Parker 1972). It recovers rapidly from slashing. It is eaten by cattle, but is less 

valuable as fodder than many of the pasture plants it is able to smother. Nevertheless, it is 

viewed by some as being useful to control soil erosion, to serve as a mulch when cut, and 

as being preferable to many alternative plants that might occupy the space vacated by its 

control. In its native habitat it is seldom a weed. 


MAJOR SPECIES 

TROPICAL AND SOUTH AMERICA 

Valuable information is available on 9 major and 22 minor natural enemies of M. micran- 

tha in its native region as a result of studies by Cock (1982a,b) and Freitas (1991). 

Several of the major natural enemies are reported to be promising biological control 

agents (Table 4.14.1) and all these, and probably some of those less extensively studied 

(Table 4.14.2), are worthy of serious consideration. Details of their biology and possible 

relevance for biological control are summarised below. 

Table 4.14.1 Major natural enemies of Mikania micrantha in its native range in 

Central and South America (after Cock 1982a). 

INSECTS 

Thysanoptera 


Liothrips rnikaniae 

Hemiptera 

TlNGlDAE 

Teleonernia sp. or spp. nr prolixa 

Coleoptera 

CHRYSOMELIDAE 

Desmogramma conjuncta 

Echorna rnarginata 

Echorna quadristillata 

Physimerus pygrnaeus 

APlONlDAE 

Apion luteirostre 

CURCULIONIDAE 

' Pseudoderelomus baridiiformis 

MITE 

ERlOPHYlDAE 

Acalitus sp.


Table 4.14.2 Additional natural enemies of Mikania micrantha, not known to be 

polyphagous, from Central and South America (Cock 1982a, Freitas 199 1 ). 

Species Distribution Feeding location Host range 

INSECTS 

Hemiptera 

LYGAEIDAE 

Xyonysius basalis 

(= X. inaequalis 

= X. sp. nr ementitus) 

MEMBRACIDAE 

Entylia sinuata 

Entylia sp. 

Trinidad, inflorescence 

Venezuela 

Colombia, stems and leaves 

Costa Rica 

Trinidad, Peru, stems and leaves 

Venezuela 

Ecuador 

Trinidad, stems and young 

Probably other Mikania 

spp.or other Asteraceae 

M. cordifolia 

M. cordifolia 

Micrutalis binaria M. vitifolia and 

Colombia leaves 

M. trinitaria 

MlRlDAE 

Pycnoderes incurvus 

TlNGlDAE 

Colombia, 

Ecuador, 

Costa Rica 

leaves 

probably specific 

Leptocysta sexnebulosa Venezuela 

Colombia. Peru 

mature leaves M. cordifolia 

Coleoptera 

CHLAMlSlDAE 

Exema complicata Trinidad, Peru, leaves 

Costa Rica, 

Colombia 

CHRYSOMELIDAE 

Longitarsus sp. 

nr amazonus 

Sceloenopla sp. 

Colombia, Peru leaves 

Trinidad leaves 

Diptera 

AGROMYZIDAE 

Calycomyza mikaniae Trinidad, leaf miner 

Colombia, 

CEClDOMYllDAE 

Neolasioptera sp. 

TEPHRlTlDAE 

Xanthaciura insecta 

Costa Rica 

Mikania spp., Chromolaena 

odorata and possibly 

other Asteraceae 

probably specific 

M. trinitaria 

Mikania spp. and possibly 

related Asteraceae 

Trinidad, Colombia flowers M. cordifolia 

Trinidad flower head 

Lepidoptera 

GELECHllDAE 

Onebala tegulella Trinidad, 

Costa Rica 

leaf roller 

Recurvaria sp. Trinidad flowers 

various Asteraceae 

M. vitifolia 




Species Distribution Feeding location Host range 

GEOMETRIDAE 

Chloropteryx sp. 

Eupithecia sp. 

LYCAENIDAE 

Thereus 

(= Thecla) palegon 

NYMPHALIDAE 

Tegosa claudina 

(= Tegosa similis) 

PTEROPHORIDAE 

Adaina bipuncta 

PY RALl DAE 

Lamprosema distincta 

TORTRICIDAE 

Lobesia (= Polychrosis) 

?carduana 

Phalonidia 

multistrigata 

Trinidad 

Trinidad 

Trinidad 

Brazil, Trinidad, 

Colombia 

Trinidad 

Trinidad, Panama, 

Costa Rica 

Trinidad 

Trinidad 

flowers 

flowers 

flowers 

leaves 

flowers 

leaf roller 

flowers 

flowers 




possibly specific 


possibly specific 



Acalitus sp. Acarina: Eriophyidae 

Feeding on the leaves by this eriophyid mite causes the formation of raised patches 

(erinea) in which the mites and their immature stages congregate. In Venezuela the patches 

mostly protrude on the undersurface of the leaf, whereas elsewhere they are mostly on 

the uppersurface, which may indicate taxonomic differences. At low mite densities the 

small number of erineum patches appear to have little effect on the growth and vigour of 

the plant. However, in dense infestations, erineum patches cover all the young leaves and 

spread into the flower heads, resulting in shortened internodes and reduced flowering. 

Plant vigour is significantly reduced. 

Eriophyid mites are usually restricted to a single plant species. Although erineum 

patches occur widely on M. micrantha, they were not seen on any other species of 

Mikania encountered in Cock's (1 982a) studies, suggesting a high degree of specificity. 

Predatory or scavenger mites occur quite commonly in and around the erineum patches. 

If field specificity trials with potted plants of closely related Asteraceae placed among 

heavily infested M. micrantha prove negative, this mite would be a promising biological 

control agent. Similar mites on Lantana in South America appear to discourage insect 

attack (K.L.S. Harley pers. comm.). 

Apion luteirostre Coleoptera: Apionidae 

Eggs of this weevil are laid in unopened host flower heads. Larvae feed initially on the 

petals, stigma and stamens and, later, destroy the developing seeds. They pupate in the 

flower head. The adults make small holes in young leaves. 

rn extensive field studies A. luteirostre larvae were not recorded from Chromolaena 

odorata, although they were found on M. micrantha and M. vitifolia, but not on M. cordifolia. 

Starvation tests using adults resulted in their feeding on five species of Mikania and 

on Bidens pilosa, but not on Chromolaena odorata.


Larvae of Apion luteirostre are attacked by the non-specific eulophid parasitoid 

Horismenus? aeneicollis and the pteromalid Zatropis sp. A number of Apion species have 

been used successfully in biological control of weeds programs (e.g. Emex australis and 

E. spinosa). Further host specificity trials are needed to evaluate the potential value of 

A. luteirostre. 

Desmogramma conjuncta Coleoptera: Chrysomelidae 

This chrysomelid beetle occurs widely, but at low density, on M. micrantha in Central 

and South America and a related species D. bigaria occurs on M. micrantha in 

Venezuela. 

Eggs are laid on the host leaves on which the larvae feed. Pupation occurs in the 

soil. No field records are available of feeding on plants other than M. micrantha and, in 

limited multiple choice tests, adults offered Bidens pilosa (cobbler's pegs), Chromolaena 

odorata and M. micrantha (all Asteraceae) attacked only the latter. No natural enemies 

have been recorded. 

The chrysomelid subfamily Chrysomelinae to which this species belongs includes 

several successful biological control agents such as the Chrysolina species on St John's 

Wort, Hypericum perforaturn angustifolium. If species of Desmogramma are specific to 

M. micrantha they may prove to have potential as biological control agents. 

Echoma (= Omoplata) marginata and E. quadristillata Coleoptera: Chrysomelidae 

Adults and larvae of these chrysomelid beetles feed openly on M. micrantha and 

M. cordifolia leaves and cause general defoliation. E. marginata is uncommon, but 

E. quadristillata is quite common around Turrialba (Costa Rica). They appear to have a 

low reproductive potential (Cock 1982a). In limited-choice tests, E. quadristillata fed on 

M. micrantha and M. cordifolia, but not on Bidens pilosa or Sonchus sp. (Asteraceae). In 

another experiment, no preference was shown between its two host species, but it would 

not feed on another species of Mikania, which was probably M. vitifolia. 

A tachinid pupal parasitoid Hyalomyodes triangulifer is known from E. marginata 

and a chalcidid pupal parasitoid Brachymeria russelli from E. quadristillata. 

Liothrips mikaniae Thysanoptera: Phlaeothripidae 

This thrips occurs in Colombia, Costa Rica, Peru, Surinam, Trinidad and Venezuela. The 

eggs are mainly laid on the undersurface of the host plant leaves or at the base of leaf 

stalks and the larvae feed there in groups. The prepupae and pupae are found among leaf 

litter beneath the plant and the adults return to the youngest leaves to feed, mate and 

oviposit. L. mikaniae has been found only on M. micrantha growing in full sunshine and 

it never occurs on plants in shady situations. This limits its potential effectiveness to sun- 

lit stands of the weed. The life cycle (egg to egg-laying adult) takes about 35 days, males 

living about 28 days, females about 35 days and laying between 21 and 11 1 eggs (Ooi et 

al. 1993). The feeding by larvae and adults on the young leaves produces small to moder- 

ate-sized lesions on the undersurface, which dry to form brown scars and these cause 

extensive distortion of the leaves as they grow. 

Laboratory studies in Trinidad (Cock 1982b) and field observations (Cock 198 1, 

1982a,b) show that L. mikaniae is restricted to the genus Mikania and most probably to

4.1 4 Mikania micrantha 131 

M. micrantha, although M. cordifolia and M. vitifolia may be fed on to a limited extent in 

the laboratory. Additional studies carried out in England by CIBC confirmed its host 

specificity before permission was obtained to introduce L. mikaniae to Malaysia. Rearing 

methods are described by Cock (1982b) and Ooi et al. (1993). 

Physimerus pygmaeus Coleoptera: Chrysomelidae 

This halticine chrysomelid is one of a group of five Physimerus species occurring on 

M. micrantha in South America. 

The larval feeding habits are unknown, but they may attack roots. The adults feed 

on young leaves and petioles, causing the dieback of growing tips, and they may be 

destructive when in high densities. This species is uncommon in Trinidad, where it is 

restricted to shady conditions, whereas in Colombia it also occurs in the open. 

Adults of I? pygmaeus have been found feeding on both M. vitifolia and M. hookeri- 

ana, in addition to M. micrantha. Field-collected adults fed on Bidens pilosa, but not on 

Chromolaena odorata. No natural enemies have been recorded. Further specificity tests 

with larvae and adults would be necessary before the potential of this species could be 

evaluated. Various halticine beetles, Longitarsus spp., show great promise for the biolog- 

ical control of ragwort Senecio jacobaea, Paterson's curse Echium plantagineum and 

common heliotrope Heliotropium europaeum. 

Pseudoderelomus baridiifomis Coleoptera: Curculionidae 

Larvae of this weevil are not known and may be root or stem gall feeders. The adult bur- 

rows into the flowers, damaging the petals, stamens and stigma and prevents seed pro- 

duction from the flowering head it occupies. When common, levels of 25% damage have 

been recorded. 

Adults of P. baridiiformis occur mainly in the flowers of M. micrantha, but have 

also been recorded from M. trinitaria and M. vitifolia. They occur rarely in the flowers of 

Chromolaena odorata and have been recorded once from Neurolaena lobata. No natural 

enemies are known. 

Although the level of damage caused may be considerable, further studies of life 

history and host specificity are required. 

Teleonemia sp. or spp. nr prolixa Hemiptera: Tingidae 

A taxonomic study of the bug genus Teleonemia (which contains more than 80 species) is 

required to enable the determination of correct identity of the one or more species of 

brown tingid bugs feeding on Mikania flowering heads. The species is not i? prolixa, 

which is highly specific to Lantana camara (Harley and Kassulke 19,759. 

The eggs are laid into the flower bracts and the nymphs and adults feed on the 

flower heads, but do not appear to cause much damage at low densities. Faeces deposited 

on the opening flowers may be sufficient to prevent seed production, particularly when 

these serve as a substrate for fungal growth. i? harleyi in Trinidad has a similar life cycle 

and feeding habits in Lantana camara flowers (Harley and Kassulke 1975). 

Adults and nymphs of Teleonemia were found by Cock (1982a) on a number of 

Mikania species (micrantha, vitifolia, trinitaria, hookeriana). Although T. prolixa has 

been recorded from Cinchona sp. (Drake and Poor 1938), Lantana camara (Monte 1939)


and Acacia riparia (Drake and Ruhoff 1965), the records for Cinchona and Acacia 

appear to be in error (Harley and Kassulke 1975). A parasite attacks the eggs of 

Teleonemia and the lygaeid Xyonysius sp. in M. micrantha flowers. Teleonemia scrupu- 

losa has been used in a number of countries to considerable effect to help in the control 

of Lantana camara. If the flower-feeding Teleonemia that attack Mikania cause similar 

effects through injection of saliva, they may cause more damage than is apparent at first 

sight. 

MINOR SPECIES 

Cock (1982a) and Freitas (1991) list a further 22 species of insects attacking M. micran- 

tha in Central and South America (Table 4.14.2). There are 7 species of Hemiptera, 3 

Coleoptera, 10 Lepidoptera and 3 Diptera. Five of these are considered at this stage of 

knowledge to be promising. 

Exema complicata Coleoptera: Chlamisidae 

Adults and larvae of this beetle are leaf feeders on Mikania spp., Chromolaena odorata 

and perhaps other Asteraceae. 

Longitarsus nr amazonus Coleoptera: Chrysomelidae 

Adults of this halticine beetle feed on leaves of M. micrantha and larvae probably on 

roots. Longitarsus species generally have a very restricted host range. 

Neolasioptera sp. Diptera: Cecidomyiidae 

The larvae of this fly feed within the flower head and scar the seed shell, but the effect of 

this damage on seed viability is unknown. This species is parasitised by a eulophid 

Tetrastichus sp. 

Sceloenopla sp. Coleoptera: Chrysomelidae 

Adults of this hispine beetle feed on leaves and larvae are leafminers on M. micrantha 

and M. trinitaria. Horismenus? aeneicollis is recorded as a larval parasitoid. Hispine bee- 

tles have proved to be very effective against Lantana camara. 

Tegosa claudina Lepidoptera: Nymphalidae 

Earlier referred to as Tegosa similis, the larvae of this butterfly are leaf feeders on both 

Mikania micrantha and M. cordifolia. Eggs are laid in clusters and larvae are gregarious, 

passing through six instars (Freitas 1991). 


MALAYSIA 

An extensive evaluation of natural enemies attacking M. micrantha was made in 

peninsular Malaysia prior to a decision to embark upon a biological control project (Teoh 

et al. 1985). Of the insects collected from or reared on the host plant, 2506 were classi- 

fied arid separated into commonly found and minor natural enemies (Table 4.14.3). 

Although numerous insects were found to attack M. micrantha, not only was the 

extent, of damage insignificant, but most of the abundant species were known pests of 

economic crops. For example, Halticus minutus and Lamprosema diemenalis are major


pests of leguminous cover crops, Homoeocerus serrifer attacks rice and Helopeltis spp. 

are serious pests of cocoa. None of the natural enemies recorded in Tropical and South 

America were found in the survey. A number of fungi were also found, of which 

Colletotrichium gloeosporioides was the most important, comprising 84% of the sam- 

ples. Other fungi included Colletotrichium spp., a non-sporulating brown fungus, 

Rhizoctonia spp., Curvularia spp. and Pestalotia spp. It was concluded that a strong case 

existed for the introduction of effective natural enemies. 

Table 4.14.3 Insects attacking Mikania micrantha in peninsular Malaysia (Teoh et al. 

1985). 

Species Effects 

MAJOR SPECIES 

Orthoptera 

ACRlDlDAE 

Acrida turrita 

Hemiptera 

APHlDlDAE 

Aphis spiraecola 

CERCOPIDAE 

Clovia conifer 

CICADELLIDAE 

Bothrogonia ferrugenea 

COREIDAE 

Homoeocerus serrifer 

Riptortus linearis 

MEMBRACIDAE 

Centrotypus flexuosus 

MlRlDAE 

Halticus minutus 

Helopeltis spp. 

Lepidoptera 

PY RALl DAE 

MINOR SPECIES 

Orthoptera 

ACRlDlDAE 

Catantops humilis 

Hemiptera 

APHlDlDAE 


CICADELLIDAE 

Nephotettix spp. 

Thysanoptera 

THRlPlDAE 

Isothrips spp. 

Microcephalothrips spp. 

holes in leaves 

wrinkled leaves 

yellow spots on leaves and stems 

brown spots on stems 

brown spots on leaves 

brown spots on leaves 

necrosis on stems 

necrotic lesions on leaves 

necrotic lesions on leaves 




Species Effects 

Parthenothrips spp. 

Thrips hawaiiensis 

Thrips tabaci 

Coleoptera 

CHRYSOMELIDAE 

Dactylispa bipartista 

COCCINELLIDAE 

Coelophora bissellata 

Epilachnu indica 

Diptera 

TEPHRlTlDAE 

Sphaeniscus atilus 

Lepidoptera 

AMATIDAE 

Amata huebneri 

PYRALIDAE 

Hellula undalis 

Liothrips mikaniae was introduced to Malaysia from Trinidad via England in 1989 

for additional host specificity trials. Difficulties were experienced initially in mass rearing, 

due to unsuitable environmental conditions (lighting, temperature, aeration) low 

plant nutritional quality and predators (spiders, ants, and particularly a predatory thrips, 

Xyloplothrips sp. which destroyed 90% of the culture). Also, there were differences in 

the M. micrantha plants used. Some were hairless and others hairy. Larvae hatching from 

eggs along stems of the latter found difficulty in moving among the trichomes and many 

failed to reach the nearest leaf. Nevertheless, in Trinidad, L. mikaniae was found breeding 

on both plant types. After passing tests with 18 Malaysian crop species, 13 161 adult 

thrips were released in 25 batches of 99 to 1400 at 5 different sites from April 1990 to 

June 1991, but no establishment resulted (Table 4.14.4). Further thrips were imported in 

January 1992 to extend the genetic base of the rearing colony. Two releases were made 

into a fenced site, 18 000 adults, together with their rearing plants in pots, in May 1992 

and 1500 adults about a month later. The pots were watered daily for two months. 

However, L. mikaniae gradually disappeared and, after eight months, none could be 

found. An ant that made its nest amongst Mikania leaves was observed to feed voraciously 

on adult, larval and pupal stages of L. mikaniae, but showed little interest in eggs, 

whereas a predatory Haplothrips sp. preferred eggs. About 20 other potential predators 

were evaluated, but were not implicated (Liau et al. 1991, 1993, Norman et al. 1992, 

Teoh et al. 1985, Ooi 1993, Ooi et al. 1993). 

SOLOMON ISLANDS 

A consignment of L. mikaniae was sent from Malaysia to the Solomon Is and released in 

the field in 1988, but the site was flooded shortly afterwards. Further releases were made, 

but no establishment has occurred (M. Vagalo pers. comm.). It was suggested that there


may be differences in hospitability to L. mikaniae of the host plant between the 

Caribbean and Solomon Is. 

PAPUA NEW GUINEA 

Part of a consignment of L. mikaniae sent to the Solomon Is in 1989 was taken to Papua 

New Guinea, but there is no information on its fate (Williams et al. 1990), although it is 

believed to have died in quarantine (F. Dori pers. comm. 1993). 

SRI LANKA AND ASSAM 

In Sri Lanka and Assam it has been found that the plant parasite Cuscuta chinensis will 

suppress Mikania and prevent it spreading into tea plantations, although C. chinensis is 

not sufficiently selective to be used in the plantations themselves (Parker 1972). On 

Espiritu Santo (Vanuatu) a related species Cuscuta campestris is reported to suppress M. 

micrantha (M.J.W. Cock pers. comm.). 

Table 4.14.4 Liberations for the biological control of Mikania micrantha. 

Species Where From When Result References 

Thysanoptera 


Lioihrips mikaniae Malaysia Trinidad via UK 1991 - Liau et al. 1991, 

1993, Norman et 

al. 1992 

Solomon Is Trinidad via UK 1988 - M. Vagalo pers. 

comm., Williams et 

al. 1990 

Comment 

Mikania micrantha is an introduced weed of widespread importance in plantation crops 

throughout Southeast Asia. It is not a significant weed in its native range in tropical 

Central and South America, where it is attacked by a wide range of arthropod natural 

enemies. Several appear to be not only damaging to the weed, but also highly specific. It 

must be concluded that M. micrantha is a highly appropriate target for an attempt at bio- 

logical control. 

The failure of Liothrips mikaniae to become established in Malaysia is disappoint- 

ing. The most probable explanation, partly supported by observations, is that heavy pre- 

dation was the cause. Although the same guild of predators may be widespread on conti- 

nental Southeast Asia, the same may not apply to island nations, particularly those in the 

southern Pacific, which have a far less diverse fauna and releases there might lead to suc- 

cessful establishment.


Mimosa in visa 

(after Holm eta/. 1977)

Map 4.15 Mimosa invisa 

4.15 Mimosa invisa 

Mimosa in visa 

Creeping sensitive plant, Mimosa invisa, is native to Tropical America, where it is not 

regarded as a weed. 

At least 70 species of insects attack it in Brazil and additional species elsewhere. 

Detailed studies have been made on two Hemiptera (Heteropsylla spinulosa and 

Scamurius sp.) and a moth (Psigida walkeri). H. spinulosa has caused extensive damage 

to M. invisa following its establishment in Australia and promising early results in Fiji, 

Papua New Guinea and Pohnpei, but disappointing results in Western Samoa. Scamurius 

sp. has failed to become established and P. walkeri is still under investigation. 

The prospects for biological control of M. invisa appear to be good, although 

additional natural enemies may have to be considered.


4.15 Mimosa invisa Martius ex Colla 

Mi mosaceae 

creeping sensitive plant; banla saet (Cambodia), borang, pis koetjing, rembete 

(Indonesia), duri semalu (Malaysia), makahiyang lalaki (Philippines), maiyaraap 

thao (Thailand), co trinh nu moc (Vietnam) 

Rating 

+++ Phil 

18 ++ Myan, Thai, Laos, Viet, Msia, Sing, Indo 

+ Camb 

Origin 

Tropical America. In Brazil southwards from Bahia to Paranh and westwards to Paraguay 

and tropical northeast Argentina; also lowlands of Central America from Veracruz 

(Mexico) southeastwards to Panama and adjacent Colombia. 

Distribution 

In addition to the above there are, in the Americas, scattered records from Brazilian 

Amazonia, the Guianas, Jamaica, Hispaniola and Cuba. M. invisa is widely distributed in 

Southeast Asia and the Pacific, also in Queensland, India, Sri Lanka, Taiwan and Nigeria. 

It was first recorded in Java in 1900 (Soerjani et al. 1987). 


Mimosa invisa is a fast growing, abundantly thorny, biennial or perennial shrub with 

angular branching stems that become woody with age. The leaves are alternate, bipinnate 

and compound. The pink to purple globular flowers are borne on a short prickly stalk 

arising from a leaf axil. The seed pods are covered with stiff bristles and separate at 

transverse grooves into two to four, single-seeded segments. 

The genus Mimosa does not occur naturally in Southeast Asia or Australia. M. 

invisa is one of three weedy species of Mimosa in this region, all of which are treated in 

this volume. They may be distinguished (i) by the number of pinnae in the leaves: M. 

invisa 4 to 9 pairs; M. pigra 6 to 16 pairs; and M. pudica 1 to 2 pairs and (ii) the size of 

the pods: M. invisa 4 seeds per pod, M. pigra 12 to 24 seeds. In addition, M. invisa stems 

have a dense covering of small prickles, whereas M. pigra stems have a sparse covering 

of large prickles (Lonsdale 1992). 

M. invisa folds its pinnate leaves when touched, but is not as sensitive as some other 

species, such as M. pudica. The leaves fold at nightfall. 

unlike the situation in the more tropical regions, such as the Philippines where 

M. invisa flowers all year round, in central and southern Brazil it only flowers from the 

end of January to mid April. Seeds mature from February to the end of May and plants 

then senesce, losing most of their leaves, although a few green leaves remain at the stem

4.15 Mimosa invisa 139 

base. For two to five months green plants are difficult to find. Senescence is not due to 

water shortage as well-watered plants in the laboratory also senesce. However, germina- 

tion occurs when moisture is available, so young plants may appear after showers of rain 

(Garcia 1982b). 

Importance 

M. invisa scrambles vigorously over other plants, forming dense tangled thickets up to 2 

m high. It is a nitrogen-fixer and its extremely rapid growth smothers useful plants and 

other weeds. Its sharp, recurved thorns make stock reluctant to graze on it and difficult 

for them to penetrate the stands. Crops infested with M. invisa are difficult to harvest 

because the thorns puncture and lacerate the hands of the workers. It is common along 

roadsides and in moist waste places. It causes major problems in coconut, tea and rubber 

plantations, sugarcane and pineapple fields, crop lands and pastures. It is not a problem 

in the Americas, western Asia, East Africa or Europe, but is a serious weed in Southeast 

Asia and the Pacific. A spineless variety, M. invisa inermis, has been suggested as a trop- 

ical pasture legume, but its tendency to revert to the thorny type and its potential toxicity 

has discouraged its use (Waterhouse and Norris 1987). 

In the Americas, M. invisa is most common in the Paranh basin in the State of San 

Paulo (Brazil), but even there pure stands are not common and it does not appear to 

invade nearby crops. In forest regions it occurs as thickets among grasses along roads, 

river banks and in waste places; it occurs more commonly on the fringe of cities (Garcia 

1982b). 


Information up to 1986 was summarised by Waterhouse and Norris (1 987). 

M. invisa is seldom troublesome in Brazil or Argentina and some 70 species of 

insects and two fungi are known to attack it in Brazil (Table 4.15.1). Additional insects are 

known in the Americas, but they have not yet been studied. A pathogenic fungus 

Corynespora cassiicola kills M. invisa in Australia (Haseler 1984), heavy infestations of 

scale insects attack it in Fiji (Mune and Parham 1967), a non-specific lymantriid larva 

feeds on young leaves and flowers in Thailand (Napompeth 1982) and a pierid butterfly 

Eurema sp. breeds on it in Papua New Guinea and New Britain (T.L. Fenner pers. comm.). 

More than half of the insects attacking M. invisa in Brazil have not yet been identi- 

fied. Indeed it is probable that most of these are undescribed species and, if so, it follows 

that there is no published information about them. Where possible, identification, even to 

a genus, may be valuable. For example, species of the genus Heteropsylla are restricted 

to legumes, with known hosts only in the Mimosaceae or Caesalpiniaceae. Of the 35 

described species with recorded hosts, 31 are specific to a single host (Hodkinson and 

White 1981, Muddiman et al. 1992). 

Only a few of the 70 insect species attacking M. invisa are known as agricultural 

pests (Table 4.15.1). From the remainder, preliminary observations on a subgroup of 

about 10 species led to detailed studies on three, Heteropsylla spinulosa, Scamurius sp. 

and Psigida (= Psylopigida) walkeri.



AUSTRALIA 

Large numbers of Heteropsylla spinulosa were released in coastal Queensland, com- 

mencing in 1988 (Table 4.15.2). This involved 33 field sites, averaging thousands of 

insects per release (M. Ablin pers. comm. 1990). Within two years, the psyllid had dis- 

persed widely into all infestations of M. invisa in pastures. Dense clumps of the weed 

were reduced to small masses of bare stems with stunted growing tips, leading to other 

vegetation reestablishing itself. Seed production from severely affected plants was 

reduced by 85 to 100% (Ablin 1992, Anon 1988). It did not attack M. pudica plants near- 

by. A more recent assessment, using insecticide-produced exclusion, found that H. spinu- 

losa reduced seed production on average by SO%, growing tip elongation by 77% and the 

growth rate of tips by 50% (Ablin 1993a). Although M. invisa may still produce clusters 

of seed pods when damage is high, the pods contain very few viable seeds. Mature plants 

support low populations of H. spinulosa during the dry season from July to November. 

Thereafter, psyllid abundance increases with the onset of summer rains, with peak num- 

bers in April or May (M. Ablin pers. comm. 1993). 

Scamurius sp. was liberated in Queensland from 1987 to 1990 and proceeded to kill 

the tips of many shoots (Anon 1988). However it did not become established. It was also 

released against Mimosa pigra in the Northern Territory where it survived for several 

months, but fecundity was very low and the colony died out (M. Ablin pers. comm. 

1993). 

FIJI 

H. spinulosa was brought in from both Western Samoa and Australia in 1993 and, after a 

generation in quarantine, was liberated in Nadi in June. Six weeks later all stages were 

seen in the field. A mealy bug and Tetranychus sp. mites are occasionally found on 

M. invisa in the field (S.N. La1 pers. comm. 1993). 


Heteropsylla spinulosa from Australia was reared through one generation in quarantine 

in Port Moresby and released early in 1993 in the Ramu Valley near Lae. Within a few 

months it had severely damaged M. invisa and killed many plants (F. Dori pers. comm. 

1993). 

POHNPEI 

Ten months after release at Palikir, Pohnpei H. spinulosa became abundant on M. invisa 

and subsequently killed many plants. Many psyllids were transferred to M. invisa in other 

areas (N.M. Esguena pers. comm. 1993). 

WESTERN SAMOA 

A total of 47000 nymphs and adults of Heteropsylla spinulosa from Australia were liber- 

ated in Western Samoa in 1988 and 1989 and, a year later, the psyllid was reported at 

some sampling sites to have reduced seed production, although not the area infested 

(Willson and Ablin 1991). 

Scamurius sp. was also liberated in Western Samoa in 1989 and was seen in the 

field after more than one generation, but not in more recent times. There have been no

4.15 Mimosa invisa 141 

reports of its effects. M. invisa continues to be a serious weed, with more than 85% of 

villages on the main island of Upolu being infested (Willson and Garcia 1992). 

Important Natural Enemies 

INSECTS 

Heteropsylla spinulosa Hemiptera: Psyllidae 

The average development period of this small (2.5 mm long) pale green, Brazilian psyllid 

is about 28 days. High populations cause severe stunting and distortion of the leaves and 

growing tips; flowering is reduced or even prevented. A sticky honeydew is produced 

which encourages a dense growth of sooty moulds. Females attach eggs by means of a 

pedicel inserted into the plant tissue between overlapping leaflets. Young nymphs live 

hidden between the leaflets, whereas adults feed on leaflets and shoots. 

In Brazil reduviid bugs pierce nymphs with their proboscis and withdraw them from 

their shelters, whereas larvae and adults of the predatory coccinellid Eriopis connexa are 

only able to capture exposed nymphs. Nymphs are also attacked by an encyrtid wasp 

Psyllaephagus yaseeni (Willson and Garcia 1992). The predatory vespid wasp 

Protonectarina sylveiriae attacks nymphs and an unidentified wasp causes up to 13% 

parasitisation (Garcia 1985). 

In extensive host specificity tests H. spinulosa adults and nymphs were unable to 

live on any plant other than M. invisa and its spineless variety M. invisa inermis. In the 

field it did not attack M. pudica, even when large infestations of M. invisa were 

destroyed and M. pudica was common nearby (M. Ablin pers. comm. 1993). Eggs were 

laid on 18 other plant species, but only under glasshouse conditions (Garcia 1985, Wild 

1987, Willson 1987, Willson and Garcia 1992) and Heteropsylla spinulosa was judged 

safe to liberate in Australia and four other countries (Table 4.15.2) 

Psygida walkeri Lepidoptera: Cercophanidae 

This moth is widespread in Brazil and Colombia. Females have a wing span of up to 5 

cm. When fully grown its greenish, spiny larvae may reach a length of 5 cm. They feed 

voraciously on leaves, flower buds, tender seed pods and on the top 30 cm of tender 

stems and branches, preventing both flowering and seed production. The life cycle takes 

about 2 months and there are 3 generations a year. There is a pupal diapause of up to 4 

months in Brazil 

Larvae of P. walkeri have been found in the field on M. invisa, M. rixosa, M. vel- 

loziana and once on M. somnians. They have not been found in the field in Brazil on 

other leguminous plants near M. invisa plants bearing larvae, nor on any plants of eco- 

nomic importance (Garcia 1983). However, under artificial conditions larvae can be 

reared on black wattle Acacia mearnsii and may also feed on Mimosa pudica and 

Leucaena leucocephala. Although adults will oviposit on A. mearnsii, no attack has been 

observed in the field (Haseler 1984). Further host specificity testing is in progress in 

Australia.


Scamurius sp. Hemiptera: Coreidae 

Both nymphs and adults of this large (up to 2.2 cm) coreid bug feed on the shoots, caus- 

ing them to collapse, thereby inhibiting vegetative growth and flowering. First instar 

nymphs moult whether fed or not and, after five instars, mature in about seven weeks. 

There are about four generations a year, from early summer to autumn, and adults may 

live for six months or more. 

Adults were found to probe many species of plants, but to feed only on species of 

Mimosa. Nymphs were able to develop on M. invisa and on two other weedy species, 

M. pigra and M. pudica, but not on other plants (Garcia 1984, Wild 1986, 1987). This 

species was approved for release in Australia and Western Samoa (Table 4.15.2). 

FUNGUS 

Corynespora cassiicola 

This stem spot fungus is very common in hot humid weather in north Queensland, Papua 

New Guinea (Keravat, Rabaul) and Western Samoa. It can be very damaging to M. invisa 

if environmental conditions are suitable (Willson and Ablin 1991). The strain involved 

appears to be specific to M. invisa, although fungi with the same specific name are 

reported from cowpea, papaya and tomato. If suitable environmental conditions persist in 

the field M. invisa plants shed their leaflets and stems die back as lesions cover the plant 

(M. Ablin pers. comm. 1993). 

Comments 

The use, in the future, of H. spinulosa from Brazil against M. invisa is complicated by 

several introductions of natural enemies that have already been made by Southeast Asian 

countries (eg. Thailand, Philippines). These introductions were of two parasitic wasps 

(Tamarixia leucaenae and Psyllaephagus yaseeni) native to the Caribbean and Central 

America (Noyes 1990) and one or more predatory coccinellids against a pest psyllid 

Heteropsylla cubana which appeared from the Americas in the 1980s. This pest can 

cause severe damage to Leucaena leucocephala which is widely planted for firewood and 

as fodder. Tests showed that these natural enemies of H. cubana would also attack 

H. spinulosa (Baker 1990). As a result, several countries where M. invisa is a serious pest 

(Australia and most Pacific islands, but not New Caledonia) have deferred introducing 

natural enemies of H. cubana. 

Several interesting points, relevant to any investigation for natural enemies in 

South America, emerge from the M. invisa project there: 

Before the studies of C.A. Garcia in the early 1980s almost nothing was known about 

the insects attacking it, yet within a year 57 insects were listed from Brazil (Garcia 

1982a,b) and within two years a further 10. No records are available from elsewhere 

in the Americas, except of a Heteropsylla sp. from Colombia (Garcia 1983). 

It has not been possible for taxonomists to assign a specific name (and sometimes not 

even a generic name) to the majority of insects collected. Some were not previously 

represented in any museum collections and many are almost certainly undescribed 

species.

4.1 5 Mimosa invisa 143 

Only one (Scamurius sp.) of the three insects eventually selected for detailed study 

was recognised in the first survey which yielded 57 species. It is clear that follow up 

surveys are essential. 

No study has yet been made of the insects attacking M. invisa over a very large area 

of its presumed native range. From brief observations in Colombia Garcia (1 983) 

commented that larvae of Lepidoptera were 'quite similar to those collected off M. 

invisa in Brazil. Coleoptera in general look different'. 

Should existing biological control be considered inadequate, further detailed studies 

covering the entire native range of M. invisa might well reveal additional, adequately 

specific insects attacking it. 

Examination of the host specificity of more of the insects recorded from Brazil (Table 

4.15.1) might also reveal further adequately specific insects. 

H. spinulosa does not thrive under either very wet or very dry conditions. Its popula- 

tions depend upon the availability of green foliage and, in the dry season, are found 

on pockets of green foliage. A flush of growth after rain leads to a build up in popula- 

tions to a level that severe damage may be caused, sufficient to kill many M. invisa 

plants. 

Table 4.15.1 Natural enemies of Mimosa invisa in Brazil (from Garcia 1982a,b, 1983 

and his unpublished monthly reports). 

Species Comment 

INSECTS 

Orthoptera 

TETTlGONl IDAE 

sp. 1 

sp. 2 

attacks flowers 

attacks leaves 

Hemiptera 

CERCOPIDAE 

Tomaspis (= Zulia) enteriana 

SP . belongs to Gyopinae 

CICADELLIDAE 

sp. I common on Mimosa invisa and M. pigra and 

colonises Acacia mearnsii and M. scabrella during 

the dry season 

sp. 2 

COREIDAE 

Scamurius sp. 1 see text 

Scamurius sp. 2 

MlRlDAE 

Horciacinus signoreti (= H. argentinus) 

Taylorilygus pallidulus 

MEMBRACIDAE 

Ceresa ustulata 

an agricultural pest 

Enchenopa gracilis 

Micrutalis sp. 





PENTATOMI DAE 

Acrosternum herbidum 

Dichelops furcatus an agricultural pest 

Edessa meditabunda an agricultural pest 

Euschistus tristigmus cribarius 

Euschistus luridus 

Piezodorus guildinii an agricultural pest 

PSYLLIDAE 

Heteropsylla spinulosa see text 

Heteropsylla sp. from Colombia 

THYREOCORIDAE 

Gyrocnemis sp. 

Coleoptera 

CHRYSOMELIDAE 

Colaspis sp. 

Cryptocephalus viridiaeneus 

?Hilax sp. 

Lactica sp. 

Lexiphanes ?semicyaneus 

Lexiphanes sp. 

Metaxyonycha pallidula 

Nodonota sp. 

Pachybrachys sp. 

System s-liftera 

Temmodachrys sp. nr aphodoides 

CURCULIONIDAE 

Asynonychus godmani 

(= Pantomorus cervinus) 

Chalcodermus sp. 

Chalcodermus sp. nr segnis 

Hypanthus sp. 

Promecops sp. 1 

Promecops sp. 2 

Sibinia aspersa 

Sibinia ?subulirostris 

Sibinia sp. 

Lepidoptera 

AMATIDAE 

SP. 

CERCOPHANIDAE 

Psigida walkeri 

GEOMETRIDAE 

sp. 1 

sp. 2 

most Chrysomelidae were present ib low numbers. 

adults eat M. invisa leaves 

excellent defoliator; also attacks M. pigra 

adults eat leaves and show strong preference for 

M. invisa, but will also attack M. pudica, Calliandra 

selloi and Acacia mearnsii 

larvae bore stems, brown adults eat leaves 

black adults bore green seeds, but did not attack pods 

of 7 other legumes 

adults eat ovaries, larvae the seeds; recorded from 

Mimosa albida and M. quadrivalis, but would not 

attack six other legumes including Leucaena 

leucocephala 

seminicola group, larvae eat seeds, adults the leaves 

and ovaries and are also found in Mimosa rixosa 

red hairy larva 

see text 

debris-covered larva 

twisted larva 


FUNGI 


sp. 3 

sp. 4 

sp. 5 

HESPERIIDAE 

Cogia (= Caicella) calchas 

LYCAENIDAE 

Hemiargus hanno 

Tmolus (= Thecla) azia 

NOCTUIDAE 

sp. 1 

sp. 2 

sp. 3 

sp. 4 

PlERlDAE 

Eurema tenella 

TORTRICIDAE 

sp. 1 

sp. 2 

sp. 3 

sp. 4 

FAMILY UNKNOWN 

Cercospora canescens 

Fusarium sp. 

Uredo mimosae-invisae 

reddish green larva 

common slim larva 

thick twig larva 

Table 4.15.2 Liberations for biological control of M. invisa. 

4.1 5 Mimosa invisa 145 

occurs in Mexico and Argentina; eggs laid also on 

M. pudica, M. scabrella, Indigofera anil, Skranquia 

sp., larvae are heavily parasitised 

larvae eat leaves, flowers and seed pods; also 

M. pudica flowers 

larvae eat leaves and flowers; also M. pudica and 

groundnut flowers 

velvet black larva; also on M. pudica, M. scabrella, 

Acacia mearnsii, Calliandra selloi, Leucaena 

leucocephala 

reddish larva 

green larva 

small green larva 

occurs in Brazil, Argentina, Paraguay, Bolivia; larvae 

defoliate M. invisa; also eat M. pudica, M. scabrella, 

Acacia mearnsii and (reluctantly) Leucaena 

leucocephala 

flowerltwig roller 

pod eater 

larvae eat leaves 

larvae roll flowers 

4 species, two bore in the stem tips, 1 eats pods 

and I the flowers 

from Venezuela (H.C. Evans pers. comm. 1992) 

Species From Liberated When Result References 

Heteropsylla spinulosa Brazil Queensland 1988 + Ablin 1992, Anon 1988 

Fiji 1993 ? S.N. Lal pers. comm. 1993 

Papua New Guinea 1993 + Ablin 1993b, 


Pohnpei 1992 + N.M. Esguerra pers. comm. 

1993 

Western Samoa 1988 + Willson & Garcia 1992 

Scamurius sp. Brazil Queensland 1987 - Anon 1988 

Western Samoa 1988 ?


Mimosa pigra 

(after CSIRO. 1992)

Map 4.16 Mimosa pigra 

4.16 Mimosa pigra 

Mimosa pigra originated in the area extending from Mexico to Amazonia and 

Venezuela. Four beetles and two moths have been established on M. pigra in Australia 

in the past 10 years. Two stem-boring moths Neurostrota gunniella and Carmenta 

mimosa are having a significant effect on the vigour of the weed. N. gunniella has 

spread widely, infests most stems and is reducing seed production. All except N. gun- 

niella have been liberated in Thailand. N. gunniella was not liberated because it can 

attack the aquatic vegetable Neptunia oleracea. However the two seed-feeding bruchids 

are destroying up to 20% of the seed in Thailand 

A specific, highly pathogenic fungus, Phloeospora mimosae-pigrae, has been 

approved for release in Australia and six insects and a rust fungus are under study. 

There are grounds for confidence that a group of natural enemies will become avail- 

able that, acting together, will cause significant damage to M. pigra.


4.16 Mimosa pigra L. 

Mimosaceae 

giant sensitive plant; mai yah raap yak, maiyarap ton, chi yop luang (Thailand); 

kembang gajah, semalu gajah (Malaysia); trinh nu nhon (Vietnam); putri malu 

(Indonesia) 

Rating 

+++ Thai 

15 ++ Myan, Laos, Msia, Sing, Indo 

+ Camb, Viet 

Origin 

Mexico, southern Venezuela, or central Amazon basin. 

Distribution 

Throughout the tropics and still spreading. It was an early invader of tropical Africa and 

is spreading aggressively in northern Australia and Southeast Asia. It is not present in the 

Philippines or the oceanic Pacific. Details of its distribution and time of recognition in 

various countries are given in Lonsdale (1992). 


M. pigra is a perennial leguminous shrub, growing up to 6 m high on a wide range of 

soils, and found in moist open sites with a rainfall between 750 and 2250 mm in the trop- 

ics. Its leaves are bipinnate and sensitive to the touch, through movements of the petiole 

and pinnules. Petioles bear a slender prickle at the junction of each of the 6 to 16 pairs of 

pinnae and sometimes have stouter prickles between each pair. The stems bear 

broad-based, sharp thorns up to 7 mm long. Mature plants have many branches growing 

from the base, with a skirt of adventitious roots forming in seasonally inundated sites. 

They have a large central taproot which penetrates 1 to 2 m deep and a lateral root system 

that extends up to 3.5 m from the stem at a depth of about 5 cm. The flowers are mauve 

to pink, massed in globular heads lcm in diameter, with each head containing about 100 

flowers. Seed pods are produced in clusters of about 7, are densely bristly, 3 to 8 cm long 

and break transversely into segments each containing a seed. The bristles facilitate float- 

ing and thus rapid spread of the weed along river systems. In regions with pronounced 

wet and dry seasons, the former is the main period of growth, with flowering mainly 

from mid to late wet season. Development from flower bud to ripe seed takes about 5 

weeks (Lonsdale 1992). Average seed production is about 9000 seeds per plant, but up to 

220000 has been recorded. Although most seeds that lodge on or near the soil surface 

probably germinate within two years many seeds deeper in the soil lie dormant for long 

periods (at least 23 years). 

Previously, two varieties were recognised var. pigra and var. berlandieri, of which 

only pigra has spread around the world. Variety berlandieri has recently been renamed 

Mimosa asperata (Barneby 1989, Lonsdale 1992).

Importance 

4.16 Mimosa pigra 149 

In tropical America M. pigra usually occurs as small clumps of multi-stemmed plants 

growing in seasonally flooded habitats. However, in many countries to which it has been 

introduced, M. pigra is a serious weed of wetlands. Dense, prickly thickets compete with 

pastures, prevent access to water and hinder mustering. The thickets exclude native vege- 

tation and so alter the environment that many native plants and animals are eliminated or 

seriously affected (Lonsdale 1992). The weed leads to sediment accumulation in irriga- 

tion systems and reservoirs and, as the seed segments float, many end up in fallow rice 

paddies where they germinate rapidly. River sand containing seeds helps to establish new 

infestations when transported to building sites, road constructions etc. 

Cattle and horses occasionally browse on young plants and some wild animals find 

it acceptable, particularly as a dry-season browse but, in general, it seems to be of low 

palatability. The leaves contain low levels of the toxic amino acid mimosine. M. pigra 

was introduced to Thailand in 1945 as a green mulch crop and for erosion control in rice 

paddy irrigation channels, but it was soon found that the problems associated with it far 

outweighed any advantages (Wara-Aswapati 1983). However it is used still as a source of 

firewood and bean poles, although it is now regarded as a very serious weed. 


Surveys for natural enemies have been made in Brazil, Mexico, Venezuela (Harley et al. 

1983), Honduras (Habeck and Passoa 1982), Costa Rica (Forno 1992) and most recently 

in Belize and Cuba (I.W. Forno pers. comm.). In Honduras more than 60 species were 

listed (2 Orthoptera, 27 Hemiptera, 1 Diptera, 15 Coleoptera and 15 Lepidoptera). 

Although a full list of insects attacking M. pigra in its native range has yet to be pub- 

lished, a diverse group of over 200 is known to occur (Forno et al. 1991b). Only 12 

species are considered to be pests of agriculture and at least 45 have habits that are likely 

to lead to restricted host specificity, such as gall forming, leaf mining or stem boring. It is 

suggested that some 10% may be adequately host specific. These are likely to attack dif- 

ferent parts of the plant causing complementary damage, so that the prospects for biolog- 

ical control appear good (Forno et al. 1989b, Wilson et al. 1990). 

Six of the tropical American species of insects have been liberated (Table 4.16.1) 

five more were eventually not released after tests showed (or suggested) that they are 

insufficiently host specific (Table 4.16.2) and a further six are currently being examined 

in Australia or Mexico for host specificity (Table 4.16.3). However, the list of potential 

insects for consideration is far from exhausted. Host testing of agents for M. pigra is dis- 

cussed by Forno and Harley (1 992). 

Two fungal pathogens of M. pigra (Diabole cubensis and Phloeospora mimosae- 

pigrae) cause considerable damage in Mexico in spite of being extensively attacked by 

hypeiparasitic fungi. Without these, the pathogens should prove even more damaging. 

Other, less damaging fungi include Colletotrichium gloeosporioides, Pestalopsis sp., 

Phomopsis sp. and Oidium sp. (Evans 1988, 1990, Evans and Seier 1991, Evans et al. 

1993).


Table 4.16.1 Releases for the biological control of Mimosapigra. 

Species Part attacked Liberated Result References 

Coleoptera 

APlONlDAE 

Coelocephalapion 

aculeatum 

BRUCHIDAE 

Acanthoscelides 

puniceus 


quadridentatus 

CHRYSOMELIDAE 

Chlamisus 

mimosae* 

Lepidoptera 

GRAClLLARllDAE 

Neurostrota 

gunniella 

flower buds 

seeds 

seeds 

pinnae & 

stems 

pinnules & 

stems 

Australia 1992 

Thailand 1991 


Thailand 1984 

Vietnam 1987 


Thailand 1984 

Vietnam 1987 


Thailand 1986 

Vietnam 1990 

SESllDAE 

Carmenta mimosa stem 


Thailand 199 1 

* Introduced from Brazil, the remaining 5 insects from Mexico. 

Fomo et al. 1994, 

Wilson et al. 1992 

Wilson et al. 1992 

Kassulke et al. 

1990, Wilson & 

Flanagan 199 1, 

Harley et al. 1985 

Julien 1992 

Fomo et al. 199 1 b, 

Harley et al. 1985, 

Kassulke et al. 1990, 

Wilson & Flanagan 

1991 

Fomo et al. 1991b, 

Harley et al. 1985 

Julien 1992 

Julien 1992 

Julien 1992 

Julien 1992 

Australia 1989 Davis et al. 1991, 

Wilson & Flanagan 

1990 

Fomo et al. 1991a, 

Julien 1992 

Host specificity tests indicate that Phloeospora mimosae-pigrae is specific to M. 

pigra and Australian authorities have granted permission to liberate this pathogen. 

Testing of Diabole cubensis is still in progress. 


AUSTRALIA 

M. pigra was probably introduced to Australia at Darwin sometime during the 20 years 

before 1891 (Miller and Lonsdale 1987, Lonsdale et al. 1989) and, after a slow start, under- 

went a population explosion in the late 1970s and, by 1992, had already covered some 800 

km2 of wetlands (Lonsdale 1992). In its century of occupation, at least 114 species of 

phytophagous insects have come to attack it (5 Orthoptera, 3 Isoptera, 49 Hemiptera, 21

4.1 6 Mimosa pigra 151 

Table 4.16.2 Insects tested against Mimosa pigra, but not released (after Forno 1992). 

Species Portion attacked 

INSECTS 

Coleoptera 

CHRYSOMELIDAE 

Cryptocephalus (= Diplacaspis) nr 

miserabilis leaves 

Diplacaspis nr prosternalis stems and leaves 

Lexiphanes guerini 

young leaves 

Syphrea bibiana 

seedlings and roots 

Lepidoptera 

GELECHllDAE 

nr Aroga leaves and stems 

Table 4.16.3 Natural enemies of Mimosa pigra under investigation (I.W. Forno pers. 

comm. 1993). 

Species Portion attacked Status 

INSECTS 

Coleoptera 

APlONlDAE 

Coelocephalapion pigrae flower buds and leaves 

CURCULIONIDAE 

Chalcodermus serripes flower buds and immature seeds 

FUNGI 

Sibinia fastigiata immature seeds 

Sibinia ochreosa flower buds 

Sibinia peruana flower buds 

Sibinia seminicola immature seeds 

Diabole cubensis leaves 

Phloeospora mimosae-pigra stems, leaves, seed pods 

awaiting approval for release 

in quarantine in Australia 


under study in Mexico 



under host testing 

approved for release 

Coleoptera and 36 Lepidoptera). Of the 11 4,47 species are seldom encountered, 39 are 

occasionally found and 28 are common. Thirty of the species are known pests of cultivat- 

ed plants and all except two are thought to be polyphagous. These two (a psyllid and a 

gelechiid moth) probably feed only on a restricted number of leguminous plants. In this 

survey no plant pathogens were recorded (Flanagan et al. 1990, Wilson et al. 1990). 

In spite of this diverse insect attack there is still an enormous difference, two orders of 

magnitude, between the bank of M. pigra seeds in the soil in Mexico (a mean of 117.5lm2) 

and Australia (a mean of 12380lm2). This is believed to reflect the differential occur- 

rence of effective natural enemies in each region (Lonsdale and Segura 1987). 

The first insects to be liberated-in 1983-for biological control of M. pigra were


two seed feeding bruchid beetles, Acanthoscelides quadridentatus and A. puniceus, both 

of which established readily (Table 4.16.1). They had previously been shown to be ade- 

quately host specific (Kassulke et al. 1990). Although these species have become wide- 

spread, on average they destroy only 0.8% of mature seed, so are not having much 

impact. Of the beetles sampled, 97.8% proved to be A. puniceus (Forno et al. 1991b, 

Wilson and Flanagan 1991, Wilson et al. 1992). A parasitoid, Dinarmus sp. 

(Pteromalidae), was reared from field-collected bruchids, but did not appear to be having 

much influence on beetle populations (C. Wilson pers. comm.). 

Next, in 1985, the stem feeding beetle Chlamisus mimosae was released and readily 

became established (Forno et al. 1991b) and in 1992193 large populations have been 

found at the Finniss R., Northern Territory and are severely damaging M. pigra stems 

(I.W. Forno pers. comm. 1 993). 

During 1989, two stem boring moths Neurostrota gunniella and Carmenta mimosa 

were released. N. gunniella established rapidly and, within a few months, was not only 

widespread near the release site but damaging a large number of stems (Forno et al. 

1991b, Wilson and Flanagan 1990). By 1993, it had spread to all M. pigra infestations 

and is associated with a naturally-occurring, exotic, die-back pathogen (Wilson 1992). 

There is a strong negative correlation between seed production and moth populations, 

suggesting that N. gunniela can reduce seed numbers by up to 60% (Anon 1992). 

Carmenta mimosa is very damaging to young plants and is spreading rapidly in the 

Finniss R. region where it is severely damaging stems (I.W. Forno pers. comm. 1993, 

Wilson 1992). 

The flower bud weevil Coelocephalapion aculeatum was liberated in 1992 and has 

become established, but its effects remain to be assessed (Forno et al. 1994, Wilson et al. 

1992). 

THAILAND 

M. pigra was introduced from Indonesia to the Chiang Mai province between 1947 and 

the early 1960s as a cover and green manure crop. When found useless for the purpose, it 

was employed for the control of irrigation ditchbank erosion, but has now become one of 

the worst and most aggressive weeds in the country. 

Napompeth (1981) reported 5 insect species attacking M. pigra but, of these, only 

the boring beetle Sagra femorata caused much damage. A further study (Napompeth 

1983) increased the number to 19 insects, but without adding any promising species. 

Both Acanthoscelides puniceus and A. quadridentatus were liberated in 1984 (Table 

4.16.1) and are now destroying between 1% and 20% of mature M. pigra seeds, which is 

significantly higher than that recorded for Australia (Forno 1992). The reasons for this 

different level of effect are not known. Chlamisus mimosae was liberated in 1986 and 

became established, but is not causing significant damage. The moth Neurostrota gun- 

niella, which is producing such spectacular damage in Australia, has not been liberated in 

Thaildnd because it has been shown to be capable of developing in the aquatic Neptunia 

oleracea, which is used as a vegetable. Two other species, the weevil Coelocephalapion 

aculeatum and the moth Carmenta mimosa were liberated in 1991 (Wilson et al. 1992), 

but there is no information on their establishment or impact.

4.1 6 Mimosa pigra 153 

MALAYSIA 

Acanthoscelides quadridentatus and A puniceus have become established at Kota Bharu 

in northern Malaysia adjacent to the region where they are established in Thailand (B. 

Napompeth pers. comm. 1993). 

MYANMAR 

Acanthoscelides quadridentatus and A puniceus have also become established in 

Myanmar along the border with Thailand (B. Napompeth pers. comm. 1993). 

INDONESIA 

M. pigra has been established in Indonesia at least since 1844 but is regarded as a less 

serious weed than in Thailand (Napompeth 1982, 1983). At least 10 insects were record- 

ed on it and, at times, causing considerable damage in the Bogor area (1 Orthoptera, 3 

Hemiptera, 1 Diptera, 2 Coleoptera and 3 Lepidoptera). Only one of these, a cerambycid 

borer Milothris irrorata was regarded as having any potential to cause important damage 

(Napompeth 1982). As there were doubts about its host specificity (Kalshoven 1981), it 

was introduced to Thailand for further study but did not survive in culture (Napompeth 

1982, 1992b). Acanthoscelides spp. have been found in M. pigra seed pods collected in 

Bogor in 1992, although there are no records of releases having been made (B. 

Napompeth pers. comm. 1993). 

VIETNAM 

Acanthoscelides puniceus and A. quadridentatus from Thailand were liberated in 1987 

and Chlamisus mimosae in 1990, but there is no information on their establishment 

(Julien 1992). 

Important natural enemies 

Acanthoscelides quadridentatus Coleoptera: Chrysomelidae 

A. puniceus 

These two species occur widely in Mexico and A. quadridentatus is also recorded from 

Texas, Nicaragua and Honduras. There are also two additional species, A. pigricola and 

A. zebratus, that are apparently specific to M. pigra seeds (Habeck and Passoa 1982). 

Eggs are laid during the day in or near crevices between pod segments. At 25°C each 

larva hatches after 10 days and tunnels through the pod into a single seed in which it 

completes its development. Pupation occurs in a cell and adults emerge by chewing a 

hole through the seed coat. On average A. quadridentatus females live 93 days and lay 65 

eggs, whereas A. puniceus females live 130 days and lay 178 eggs (Kassulke et al. 1990). 

Carmenta mimosa Lepidoptera: Sesiidae 

This species is native to Mexico and Cuba. In Mexico the larvae tunnel in the stems of 

M. pigra, thereby weakening the plant. The upper two-thirds of the stems frequently snap 

off, resulting in spectacular damage. Females lay 1 to 5 eggs at a time (up to a total of 

about 260) in the axils of the topmost, fully expanded leaves on a stem. At 25°C larvae 

hatch after 11 days and tunnel into the stem at a node or the swelling at the base of a leaf 

petiole. They feed on the outer layers of the plant, sometimes ringbarking it, or tunnel

154 


through the pith; they are cannibalistic if they meet another larva. They eject frass onto 

the surface of the stem. Occasionally larvae leave the stem and re-enter below the soil 

surface, grazing on and in the root, causing damage which sometimes kills the plant. 

Depending upon plant quality, there are 8 or 9 larval instars in 40 to 99 days before lar- 

vae spin a silken cocoon in which they pupate. The duration of the pupal period is 18 to 

21 days, giving an average life cycle of 98 days. Larvae can be reared on an artificial 

diet. In host specificity tests, C. mimosa was found to complete its development only on 

M. pigra (Forno et al. 1991 a, 1994, Smith and Wilson 1992, Wilson et al. 1992). 

Chalcodermus serripes Coleoptera: Curculionidae 

This weevil is native to Mexico. Adults feed on young leaves, flower buds and seeds. 

Eggs are inserted into the ventral side of pods so that they are at the embryo end of a 

developing seed. They hatch in 6 days and larvae feed on the soft developing seed, 

destroying the embryo (I.W. Forno, pers. comm.). 

Chlamisus mimosae Coleoptera: Chrysomelidae 

This species is native to Brazil. Females mate 2 weeks after emergence and then begin to 

lay eggs. Each egg is enveloped in faecal material and attached to the underside of a leaf 

by a fine stalk. At 25°C larvae hatch after 3 weeks and construct a conical case which is 

added to as the larva grows. Larval development time is 83 days and the pupal stage lasts 

25 days. Adults live up to 95 days. Adults and larvae graze on the epidermis of the grow- 

ing tips, on green stems and on leaves. In Darwin (Northern Territory) cultures of C. 

mimosae were attacked by a pupal parasitoid and predatory mites (Wilson et al. 1992). 

Coelocephalapion aculeatum Coleoptera: Apionidae 

This species is native to Mexico. It lays one egg at a time into a separate flower bud (of 

which an inflorescence contains up to 100). Larvae hatch after 2 days and feed on the 

developing flower buds, destroying the reproductive parts and sometimes the pedicel. 

Larval development takes about 7 days and pupal development 3 days. Preoviposition is 

about 7 days, adults live at least 3 months and may lay up to 5 eggs per day. Adults chew 

into the unopened flower buds and feed on the anthers and the pistil (Heard 1992, Wilson 

et al. 1992). This species could develop satisfactorily only on M, pigra (Forno et al. 

1 994). 

Coelocephalapion spretissimum and C. pigrae Coleoptera: Apionidae 

The life cycle of these species is similar to that of C. aculeatum. Adults feed on young 

leaves as well as on flower buds. The host testing of C. pigrae has been completed. 

Neurostrota gunnielh Lepidoptera: Gracillariidae 

This species is widespread in tropical or subtropical, moderately wet to semi-arid habitats 

wherever M. pigra occurs from southern Texas to Costa Rica and Cuba. It has been 

established in Australia. 

Eggs are laid singly on the ventral side of the first or second leaf from the branch tip 

and hatch about 4 days later. The first and second instar larvae are adapted for mining by 

being flattened dorso-ventrally, having large blade-like mandibles and no thoracic legs.

4.16 Mimosa pigra 155 

Each mines up to 5 leaf pinnules. Third instar larvae are cylindrical, enter the leaf rachis 

and tunnel to the stem tip. Later instars usually tunnel down the stem. They sometimes 

leave the stem and re-enter it at a node or near a prickle. Frass is usually visible when a 

larva is inside a stem. Pupation occurs in a cocoon spun between pinnules or inside the 

stem. The outside of the cocoon is ornamented with small, pearly-white, frothy balls dis- 

charged from the anus. 

At 25°C the time from egg to adult is about 30 days and equal numbers of males 

and females are produced. Females mate on the night of emergence and lay an average of 

86 eggs, most on the second night. N. gunniella caused very severe damage to M. pigra 

in quarantine trials which demonstrated that, except for attack on Neptunia species, it 

was specific to M. pigra. The aquatic Neptunia oleracea, which is an important vegetable 

in Thailand was attacked, so it has not been released there. In Mexico N. gunniella larvae 

are heavily attacked by parasitoids (Davis et al. 1991, Forno et al. 1989a, 199 1 b). 

Sibinia spp. Coleoptera: Curculionidae 

S. fastigiata occurs from Mexico to Brazil and Peru, whereas S. seminicola occurs from 

Texas and Mexico to southeastern Brazil and northeastern Argentina. These two Sibinia 

species are larger than the two that follow. Larvae of both species develop in the pods of 

M. pigra and feed on the seeds. Larvae of S. seminicola feed on green, immature seeds 

and pupation occurs within the pods while they are still attached to the plants (Clark 

1984). 

Sibinia ochreosa occurs in Texas, Mexico, Honduras, Nicaragua, Brazil, Argentina 

and S. peruana occurs in Mexico, Guatemala, Honduras, Costa Rica, Panama, Brazil, 

Peru, Bolivia and Argentina. Larvae of these two species develop in the flower buds of 

M. pigra and, at least the former species, pupates in the flower head (Clark 1984). 

FUNGI 

Diabole cubensis Fungi: Uredinales 

This rust attacks the leaves of M. pigra in Mexico and Cuba. It is particularly common 

and damaging during the dry season when there are high day temperatures and a signifi- 

cant drop at night leading to dew formation. Five hyperparasitic fungi are consistently 

encountered, often completely overgrowing the rust (Evans 1988, 1990). 

Phloeospora mimosae-pigrae Fungi: Coelomycetes 

This fungus causes extensive defoliation during the wet season, particularly in the Gulf 

coast of Mexico, but also occurs in Trinidad, Venezuela, Colombia and Brazil. It attacks 

stems, leaves and seed pods. It is host specific to M. pigra (Evans 1988, 1990, Seier and 

Evans 1993). 

Comment 

The majority of natural enemies of M. pigra so far studied are flower or seed attacking 

insects and there are indications already that cbnsiderable amounts of seed are being 

destroyed-up to 60% from Neurostrota alone in Australia and up to 20% by bruchids in

156 


Thailand. However, starting from a seed bank of 9000 per m2, this degree of reduction is 

nowhere like limiting. 

The inhibition to tip growth produced in Australia by Neurostrota gunniella is likely 

to be far more significant and will become even more so if the borer, Carmenta mimosa, 

becomes abundant enough to weaken a considerable proportion of larger stems. 

If (when) the two apparently specific fungi are approved for release, it is confidently 

expected that they will make a major contribution, Phloeospora mimosa-pigrae in the 

wet season and the rust Diabole cubensis in the dry. 

It is highly probable that a complex of natural enemies will be required to bring 

about an adequate reduction in competitiveness of M. pigra. It is still unclear whether 

those available or under investigation will be adequate but, if required, there are addition- 

al species that could be examined and future priority might well be given to leaf, stem or 

root feeding species.

158 


Mimosa pudica 


Map 4.17 Mimosa pudica 

4.17 Mimosa pudica 

Mimosa pudica 

Very little is known about the natural enemies of Mimosa pudica in its centre of origin in 

Central America. Most of the species recorded from there or elsewhere are widely 

polyphagous and few show promise as biological control agents. Since at least three 

forms of the weed are known it would be desirable to establish which forms are weedy in 

Southeast Asia so as to enable any searches in Central America to concentrate on that 

form. On general grounds, it is probable that useful species do exist.


4.17 Mimosa pudica L. 

Mimosaceae 

common sensitive plant; paklab, sampeas (Cambodia), daoen kaget kaget 

(Indonesia) mala malu (Malaysia), makahiya (Philippines), mai yarap (Thailand) 

mac co (Vietnam) 

Rating 

+++ Myan, Sing, Indo 

17 ++ Msia, Brun, Phil 

+ Thai, Viet 

. Laos, Camb 

Origin 


Distribution 

M. pudica is widespread in tropical, subtropical and temperate areas of the world. Its dis- 

tribution and other relevant aspects were summarised by Waterhouse and Norris (1987). 

There are at least three distinct varieties (Brenan 1959). M. pudica hispida is uncommon 

in the Americas, but is established in the Philippines (Bameby 1989), the Caroline and 

Mariana Is, Queensland, India and in African savanna country. M. pudica unijuga occurs 

in Hawaii and probably in other Pacific countries where it is a major weed. 


M. pudica is low, much branched, generally perennial, slightly woody at the base, from 

15 to 100 cm high and has either an upright or a low trailing habit. Its stiff reddish-brown 

or purple stems bear scattered thorns. The hairy leaves are alternate, bipinnate and com- 

pound. They are sensitive to the touch, the petiole dropping and the leaflets being rapidly 

drawn back and folded. The pink flowers form small globular heads, each borne on a 

short hairy stalk arising from a leaf axil. Seeds are produced in pods which split into sin- 

gle-seeded segments bearing bristles, which aid dispersal by animals. In tropical coun- 

tries the weed flowers all year and each plant may produce up to 700 seeds. 

Importance 

M. pudica is a weed in 22 crops in 38 countries (Holm et al. 1977). It is common in waste 

land and is also a weed of lawns, crops, pastures and roadsides. In Southeast Asia and the 

Pacific it is a serious weed in maize, sorghum, sugarcane, tea, soybeans, upland rice, 

pinea~jples and cotton. Because of its tolerance to shading it is an important weed in plantation 

crops, such as rubber, coconuts, bananas, papaya, coffee, oil palm and citrus. In 

tropical pastures its dense growth and thorns often deter animals from feeding on suitable 

forage mingled with it (Holm et al. 1977). The thorns deter hand weeding and, as it sur-

4.1 7 Mimosa pudica 161 

vives mowing, it is a very unwelcome component of lawns. Attempts to select thornless 

types as pasture plants have not been successful. 


Some information is summarised by Waterhouse and Norris (1987). It is interesting that 

Holm et al. (1977) report M. pudica to be a widespread weed in the Caribbean, but far 

less important to the north and south of this region. This suggests that it evolved else- 

where in the Americas and has not been accompanied into the Caribbean by its full suite 

of natural enemies. Nevertheless a preliminary survey in Trinidad (Table 4.17.1) revealed 

14 insects attacking it, but they are probably polyphagous, with the possible exception of 

the arctiid caterpillar Lophocampa catenulata and the beetle, Chlarnisus sp. (Yaseen 

197 1, 1972). Perez et al. (1 988) found that the race Jilenus of Hemiargus hanno in Cuba 

appears not to attack plants other than M. pudica, although partially grown larvae of the 

Trinidad race were able to complete their development on Aeschynomene sensitiva and 

Cajanus cajan (Yaseen 1972). H. hanno Jilenus feeds readily on M. pudica seeds and is 

particularly active in spring when the weed is producing most seed (Perez et al. 1988). 

Four additional insects are known from Brazil (Garcia 1982a,b, 1983) but, so far, no 

species of Heteropsylla, although a special search for one might be rewarding. Although 

M. pudica was often encountered in surveys carried out in Mexico and Venezuela for nat- 

ural enemies of M. pigra, casual observation did not suggest that it was heavily attacked, 

less so indeed than M. invisa (I.W. Forno pers. comm. 1993). 

It is not known whether there is any differential attack by natural enemies on any of 

the three or more varieties of M. pudica, which have been established on morphological 

differences alone. 

Table 4.17.1 Natural enemies of Mimosa pudica. 

Species Country Portion of plant attacked References 

INSECTS 

Hemiptera 

COCCIDAE 

Coccus longulus Fiji 

CYDNIDAE 

Microporus 

(= Microcompsus) sp. Trinidad 

DlASPlDlDAE 

Hemiberlesia lataniae Fiji 

Pinnaspis strachani Fiji 

MARGARODIDAE 

Icerya seychellarurn Fiji 

SCUTELLERIDAE 

2 species Trinidad 

Coleoptera 

CHRYSOMELIDAE 

Chlarnisus sp. Trinidad 

stems Hinckley 1963 

stems 

stems 

stems 

flowers 

Yaseen 1972 

Hinckley 1963 

Hinckley 1963 

Hinckley 1963 

Yaseen 197 1 

flower buds Yaseen 197 1, 1972 




Species 

CURCULIONIDAE 

Chalcodermus sp. 

Promecops 

?campanulicollis 

Country Portion of plant attacked References 

Brazil 

Brazil 

seed pods 

leaves; mainly on 

M. invisa, see table 

4.15.1 

C.A. Garcia pers. comm. 

Garcia 1982a,b, 1983 

An unidentified sp. Trinidad Yaseen 197 1 

Lepidoptera 

ARCTllDAE 

Lophocampa catenulala 

GEOMETRIDAE 

Ptychamalia perlata 

GRACllARllDAE 

Neurostrota gunniella 

HESPERIIDAE 

Cogia (= Caicella) 

calchas (= ? 

Nisoniades bessus) 

?Slaphylus mazans 

LYCAENIDAE 

Calephelis sp. 

Hemiargus hanno 

Trinidad leaves Yaseen 1971 

Trinidad leaves 

Mexico leaves, stems 

Trinidad leaves, flowers 

Yaseen 1972 

Davis et al. 1991 

Cock 1985, Yaseen 1972 

Trinidad leaves, flowers Yaseen 1972 

Trinidad 

Cuba, 

Trinidad 

leaves 

leaves, flowers, pods 

also on flowers of 

Aeschynomene 

semitiva and Cajanus 

cajan (= C. indicus) 

Tmolus azia Trinidad leaves, flowers Yaseen 1972 

NOCTUI DAE 

Spodoptera litura Fiji 

Sp. 1 (velvet black larva) Brazil 

PlERlDAE 

Eurema lisa Cuba 

Eurema tenella Brazil 

also on Mimosa pudica 

and groundnut flowers 

leaves 

leaves; also on 

M. invisa 

leaves: mainly on 

M. invisa, see table 

4.15.1 

Yaseen 1971 

Dethier 1940, Perez et al. 

1988, Yaseen 1972 

Hinckley 1963 

Garcia 1982a,b, 1983 

Dethier 1940 

Garcia 1982a,b, 1983 

TORTRICIDAE 

Platynota rostrana Trinidad leaves Yaseen 1972 

NEMATODE 

Meloidogyne sp. Cuba Holm et al. 1977, 

Izquierdo et al. 1987 

FUNGI 

VIRUS 

Oidium sp. Mexico 

?Puccinia sp. Mexico 

mildew on leaves 

rust on leaves 

Evans 1987 

Evans 1987 

unspecified Germany Umrath et al. 1979




Map 4.18 Monochoria vaginalis 

4.1 8 Monochoria vaginalis 165 


Monochoria vaginalis appears to be a major weed only in Southeast Asia and then only 

in rice. Almost nothing is known of its natural enemies in India and Africa where it 

occurs, but is not regarded as important. This suggests that a survey in these regions 

might reveal promising biological control agents.


4.18 Monochoria vaginalis (Burm. f.) Presl 


monochoria; ka kiad chrach (Cambodia), phak kbiat (Thailand), rau mac la thon 

(Vietnam), etjeng padi (Indonesia), biga bigaan (Philippines), kelayar, echeng 

padi (Malaysia) 

Rating 

+++ Laos, Camb, Msia, Brun, Indo, Phil 

26 ++ Myan, Thai, Viet, Sing 

Origin 

Tropical Asia and Africa (Holm et al. 1977), but not a pest in Africa or Asia (Soerjani et 

al. 1987), although it is clearly very important in Southeast Asia. 

Distribution 

Africa, India, China, Korea, Japan, Southeast Asia to northern Australia, Fiji and Hawaii. 


A smooth, fleshy, semi-aquatic annual or perennial, 0.1 to 0.5 m tall; the plant roots in 

mud and its upper portions grow above water; stemless, base of leaves heartshaped or 

rounded, shiny deep green; petioles soft, hollow; inflorescence with a large bract and 

arising about two thirds of the way up the petiole from the base and opposite the leaf; 3 

to 25 violet or lilac flowers producing numerous, small seeds throughout most of the 

year. Seed germination and seedling growth not reduced by submergence. Old plants 

often form large clumps. 

Importance 

M. vaginalis occurs in marshy places, freshwater pools, mudflats, ditches, along canal 

banks and in rice fields. It is a very serious weed of rice in eastern and southern Asia. It 

is predominantly an annual in flooded ricefields, dying when the fields dry out, but 

developing again later from seed. 

In Taiwan, M. vaginalis produced more fresh tissue weight than any other weed in 

rice, twice that of second ranking Echinochloa crus-galli (Lin 1968). However in the 

Philippines it was outcompeted in rice by E. crus-galli (Lubigan and Vega 1971). Only in 

rice is it reported as a very widespread and important weed, except for its occurrence in 

taro in Hawaii. 

Its leaves are eaten as a pot herb in India and several parts of it are used as herbal 

medicine, the juice being prescribed for various conditions and the roots for stomach and 

liver ailments and toothache (Burkill 1935, Soerjani et al. 1987).


4.1 8 Monochoria vaginalis 167 

So little is recorded about the natural enemies of M. vaginalis (Table 4.18.1) that it is not 

possible to assess the prospects for biological control. However, the fact that it is appar- 

ently not a weed in Africa or western Asia suggests that it would be well worthwhile 

investigating these regions for suitable agents. 

Table 4.18.1 Natural enemies of Monochoria vaginalis. 


INSECTS 

Orthoptera 

ACRlDlDAE 

Gesonula punctifrons India 

Hemiptera 

CICADELLIDAE 

Macrosteles fasciffons USA rice 

DELPHACIDAE 

Tarophagus proserpina Philippines taro, cassava, sweet potato 

Lepidoptera 

NOCTUIDAE 

Spodoptera litura India widely polyphagous 

PYRALIDAE 

Elophila responsalis Indonesia Marsilea minuta, Pistia 

stratiotes, Salvinia molesta, 

S. cucullata 

Nymphulafregonalis India probably polyphagous 

SPHlNGlDAE 

Hippotion echeclus India polyphagous 

NEMATODES 

Hirschmaniella spp. rice, sugarcane, 

many weeds 

Meloidogyne graminicola rice, many weeds 

VIRUSES 

rice ragged stunt Thailand, rice 

Philippines 

Pistia virus India 

FUNGI 

Cercospora sp. India 

Doassansia sp. India 

Rhizoctonia solani India rice, potato 

Thanatephorus cucumeris Philippines rice 

a leaf blight Philippines Sphenoclea zeylanica 

CRUSTACEA 

Triops cancriformis Japan Veronica peregrina and 

some other weeds 

Sankaran & Rao 1972 

Way et al. 1983 

Duatin & Pedro 1986 



Mangodihardjo 1975 



Luc et al. 1990 


Parejarearn et al. 1988, 

Salamat et al. 1987 




Gokulapalan & Nair 1983 

Moody et al. 1987, 

Mew et al. 1980 

Bayot et a]. 1992 

Igarashi 1985


Nep hrolepis biserra ta 

(after Barnes and Chan, 1990)

Map 4.19 Nephrolepis biserrata 

4.19 Nephrolepis biserrata 169 


N. biserrata is a widespread fern that probably originated in Tropical Africa. Almost 

nothing was learnt concerning natural enemies from a literature search. A survey in its 

area of origin would be required to evaluate whether there were any natural enemies that 

might be of value for biological control.


4.19 Nephrolepis biserrata (Sw.) Schott 

Nephrolepidaceae (formerly in the Davalliaceae) 

broad sword fern; paku larat (Malaysia) 

Rating 

+++ Sing, Brun 

10 ++ Msia, Phil 

Thailand 

Origin 

Probably Tropical Africa. 

Distribution 

Pantropical. N. biserrata occurs throughout tropical Africa, ranging in the west from 

Guinea to Angola and in the east from Sudan to Durban (South Africa). It is mainly 

coastal, but infrequent in the interior. It is most abundant up to 350 m (Jacobsen 1983). In 

addition to Southeast Asia, it is also known from India, Australia, Japan, the Pacific, 

USA and Mexico. 


N. biserrata is a perennial, terrestrial or epiphytic fern. The rhizome bears abundant 

scales and produces many long stolons. The fronds are tufted, suberect to arching and 

green when young, turning brown when old. The pinnae are commonly 15 to 20 cm 

wide, exceptionally more than 30 cm. Fertile pinnae are narrower than the sterile pinnae 

and bear sori well clear of the edge. The veins are indistinct and fork once or twice. 

When rooted in the soil, the fern commonly reaches a height of 2m and, exceptionally in 

South Africa, up to 4 m. 

Importance 

N. biserrata is very common in shaded places in the lowlands wherever the conditions 

are not too dry. It can form dense masses in rubber and oil palm plantations and in 

orchards and often occurs as an epiphyte on palms. Where pineapples had been grown 

for up to 10 years in Malaysia, 90% of the viable seeds and spores in the top 15 cm of 

soil were spores of N. biserrata (in particular) and 8 other ferns (Wee 1974), enabling its 

rapid reappearance after cultivation. 


The only records of natural enemies encountered were those of an eriophyid mite on 

N. biserrata in Fiji (Mani and Jayaraman (1987) and of the nematode Aphelenchoides 

fragariae in Hawaii, but there is no evidence that a careful search has ever been made.


There have been none. 

Comment 

4.19 Nephrolepis biserrata 171 

The genus Nephrolepis contains about 20 species (Tryon and Tryon 1982), or 35 species 

(Jacobsen 1983). It is primitive among the group of oleandroid, davallioid and nephrole- 

poid ferns to which it belongs and N. biserrata and its close allies represents the more 

primitive element in the genus Nephrolepis (Nayar and Bajpai 1976). A survey for natur- 

al enemies, particularly in the areas in Africa where it occurs, would reveal whether there 

are any that might be of value in biological control.




Map 4.20 Panicum repens 

4.20 Panicum repens 


P. repens is a major grassy weed in Southeast Asia. It is probably of tropical 

AfricanJMediterranean origin, a region where it is not reported to be a problem. Very few 

natural enemies have been recorded and, without a preliminary survey in its area of ori- 

gin, it would not be possible to evaluate the prospects for its biological control.


4.20 Panicum repens L. 

Poaceae 

torpedo grass, creeping panic grass; myet kha (Myanmar) yah chan ah kat, yah 

chanagard (Thailand), chhlong (Cambodia), co 6ng (Vietnam), keruong padi, 

rumput jae jae (Malaysia and Indonesia), luya luyahan (Philippines) 

Rating 

+++ Viet, Indo 

16 ++ Thai, Msia, Sing, Phil 

+ Myan, Brun 

Laos. Camb 

Origin 

Tropical and North Africa, Mediterranean (sometimes said to be native to Asia). 

Distribution 

Panicum repens is widely distributed in the tropics and subtropics. It was introduced to 

Java about 1850, but is said not to occur in the Moluccas. No seeds are produced in 

Indonesia (Soerjani et al. 1987). 

Description 

Panicum repens is an erect, wiry, creeping, perennial grass, rooting at hairless nodes and 

bearing flowering stalks 30 to 90 cm tall. It spreads widely (up to 7 m), but does not form 

dense clumps. Its smooth, sharp-pointed, branched rhizomes are often swollen or knotty 

and have brownish or whitish scales. Its leaves are alternate. The inflorescence is an open 

terminal panicle, 6 to 20 cm long, with many tender branches pointing obliquely 

upwards. The spikelets are pale green or pale yellow and often tinged with purple. 

Importance 

Panicum repens is one of the most serious grass weeds because of its rapid rate of spread 

and the persistence and hardiness of its coarse, enlarged rhizomes. It suppresses other 

plants by its allelopathic (inhibiting) products (Perera et al. 1989). It is primarily a weed 

of moist, coastal, sandy soils, although it also grows in heavy upland soils (to 2000 m in 

Indonesia). It thrives in open sunny situations, but can stand partial shade and its rhi- 

zomes survive even prolonged dry periods. It tolerates temporary flooding, sometimes 

forms floating mats, and encroaches upon ditches, drains and watercourses. It is common 

in cultivated lands, grasslands, roadsides and gardens and is frequently reported as a weed 

in lawns. In Malaysia it is a serious weed of cocoa, coconuts and rubber, in Indonesia of 

rainfed and upland rice, cocoa, coconut, maize, rubber, sugarcane and tea and in Thailand 

of rice and orchard crops. In improved pastures it chokes out more nutritionally valuable

4.20 Panicum repens 175 

grasses. Deep ploughing favours its spread by breaking up and dispersing its rhizomes. 

P. repens is quite palatable to stock when young and has the advantage of being able 

to stand heavy grazing and trampling. However, it contains only 3.3% crude protein and 

up to 39% crude fibre, so there are other far more nutritious grasses suitable for the same 

environment (Holm et al. 1977). 


As can be seen in table 4.20.1, very few natural enemies were revealed by an extensive 

literature search. The only species not known to be both polyphagous and a pest is the 

mite Parasteneotarsonemus panici, recorded so far only from Tamil Nadu, India, 

where it was found causing rusting symptoms beneath the leaf sheath of P. repens 

(Mohanasundaram 1984). 

Absence of attack almost certainly does not represent the true situation, but rather 

that no relevant surveys have been carried out. For example, there has been little advance 

in knowledge of cecidomyiid gall flies attacking Panicum spp. since the summary by 

Barnes (1954b) in which were listed at least 13 species (Table 4.20.2). Two of these (the 

widespread rice stem gall midge Orseolia (= Pachydiplosis) oryzae and Contarinia 

(= Stenodiplosis) panici are known to be pests (Hegdekatti 1927). The scarcity of infor- 

mation from Africa points to an important gap in our knowledge and the absence of 

records from the Mediterranean, where most plants have been studied in some detail for 

native-insects, suggests that P. repens is not native to that region. 

Comment 

Clearly insufficient information is known about the natural enemies of P. repens 

(although it doubtless must have many) to provide any meaningful assessment of the 

prospects for its biological control. However, the genus Panicum contains a number of 

good to very good fodder species (e.g. P. antidotale (blue panic), P. bulbosum (bulbous 

panic), P. capillare (witchgrass), P. maximum (guinea grass), P. paludosum (swamp 

panic)) as well as several that are weedy and may cause photosensitivity or poisoning 

(e.g. P. coloratum (coolah grass), P. luzonense, P. miliaceum (millet panic, or proso), P. 

novemnerve). There are thus likely to be considerable problems in discovering organisms 

of adequate specificity. Nevertheless, P. repens is not reported as a weed of crops in trop- 

ical Africa or the Mediterranean (Holm et al. 1977) and a preliminary survey there might 

well reveal promising natural enemies.


Table 4.20.1 Natural enemies of Panicum repens. 

Natural enemies Recorded from References 

INSECTS 

Hemiptera 

CICADELLIDAE 

Thaia oryzivora Thailand 

DELPHACIDAE 

Delphacodes idonea USA 

Sogatella kolophon Australia, Pacific Is, SE Asia, 

USA, Central & S. America, 

W. Africa 

Lepidoptera 

EUPTEROTIDAE 

Nisaga simplex India 

NOCTUl DAE 

Sesamia cretica Egypt 

PYRALI DAE 

Cnaphalocrocis medinalis Philippines 

Cnaphalocrocis (= Marasmia) Philippines 

patnalis 

Parapoynx stagnalis 

(= Nymphula depunctalis) India 

MITES 

TARSONEMIDAE 

Parasteneotarsonemus panici India 

FUNGI 

Claviceps sp. 

Pyricularia sp. 

Pyricularia oryzae 

Sporisorium overeemi 

NEMATODE 

Meloidogyne graminicola 

India 

India 

Leeuwangh & Leuamsang 1967 

Ballou et al. 1987 

Ballou et al. 1987 

Patnaik et al. 1987 

Ahmed 1980 

Abenes & Khan 1990 

Abenes & Khan 1990 

Pillai & Nair 1979 

Mohanasundaram 1984 

Janardhanan et al. 1982 

Hilda & Suranarayanan 1976, 


Paje et al. 1964 

Rifai 1980 

Luc et al. 1990

4.20 Panicum repens 177 

Table 4.20.2 Gall flies (Cecidomyiidae) reported attacking Panicum spp. (after 

Barnes 1954b). 

Species Recorded from Location 

Contarinia (= Srenodiplosis) panici 

Lasioptera (= Dyodiplosis)fluitans 

Lasioptera inustorum 

Lasioptera kanni 

Lasioptera panici 

Lasioptera paniculi 

Orseolia cynodontis 

Orseolia (= Courteia) graminis 

Orseolia (= Dyodiplosis) andropoginis 

Orseolia (= Dyodiplosis) fluvialis 

Orseolia (= Pachydiplosis) oryzae 

Parallelodiplosis javanica 

Parallelodiplosis spp. 

Yugoslavia, USSR 

Panicum fluitans S. India 

USA 

S. India 

USA 

Philippines 

France, Italy, Algeria, Eritrea, 

Senegal 

Java, Sri Lanka, S. India 

Panicum jluitans S. India 

S and SE Asia, W. Africa 

Panicum indicum Sri Lanka 

Middle East, Java, Peru


Paspalurn conjuga turn 


Map 4.21 Paspalurn conjugaturn 

4.2 1 Paspalurn conjugaturn 179 

Paspalurn conjugaturn 

P. conjugaturn is of Tropical American origin, but it is recorded as an important weed in a 

number of situations in the Caribbean area. Very few natural enemies have been reported. 

A preliminary survey would be required in its centre of origin before the prospects for its 

biological control could be evaluated.


4.21 Paspalurn conjugaturn Bergius 

Poaceae 

sourgrass; paitan, rumput canggah, rumput pait (Indonesian), rumput kerbau, 

jampang canggah, buffalo grass (Malaysia), hulape (Philippines), ya hep 

(Thailand) 

Rating 

+++ Viet, Msia, Indo 

15 ++ Sing, Brunei, Phil 

Myan, Thai, Laos, Camb 

Origin 


Distribution 

Paspalurn conjugaturn occurs as a troublesome weed in Central America, West Africa 

and the islands and peninsulas of Southeast Asia and the Pacific. These are, for the most 

part, the humid tropics (Holm et al. 1977). 


P. conjugaturn is a creeping, stoloniferous, perennial grass. The stolons are up to 2 m in 

length, often reddish purple in colour and bear roots and a tuft of green to purple leaves 

at each node. 

The flower stalks are erect, range up to 60 cm, and have smooth nodes. At the apex 

of each stalk there are two racemes (flower spikes) 4 to 15 cm long. The stigmas are 

white and the anthers bright yellow. The weed can be recognised when in flower by the 

typical T-shaped inflorescence. 

Importance 

P. conjugaturn is mainly a weed of the warm, wet lowlands, although in Hawaii and Sri 

Lanka it grows up to 1875 m. It is found in waste areas and along paths and streams, its 

inflorescences trailing in the water. It is common in cultivated fields and in natural and 

poorly managed pastures and particularly in perennial or plantation crops where the soil 

is not ploughed frequently. It spreads rapidly by its stolons, forming dense masses which 

can suppress or eliminate tree seedlings and other small plants. It tolerates some shade 

and can grow on poor and acid soils. 

In the Philippines it flowers all year round and one plant can produce 1500 seeds. It 

is also dispersed by broken pieces of stolons rooting after being spread by machines used 

for cultivation. In the Philippines it is particularly important in bananas, coffee, papaya, 

rice and pineapple; in Cambodia in rice; in Malaysia in citrus, coconuts, oil palm, rice

4.21 Paspalurn conjugaturn 181 

and rubber; in Indonesia in tea, oil palm and rubber; and elsewhere in cassava, cocoa, 

lawns, maize, pastures, sugarcane and vegetables. 

P. conjugaturn is suitable for grazing only when young and the seeds of older plants 

have been reported to choke animals by sticking in their throats (Holm et al. 1977). 


The natural enemies of P. conjugatum that have been recorded in the literature (Table 

4.21.1) are almost all polyphagous and many of them are of economic importance. One 

possible exception is the bagworm moth Brachycyttarus griseus, which was originally 

described from Vietnam and is also recorded from Malaysia and the Philippines, as well 

as from Guam and Hawaii to which it has spread. It feeds on P. conjugaturn in Hawaii 

and on the grass Zoysia pungens in Guam: it probably also feeds on other grasses. In 

Guam it is parasitised by the tachinid fly, Stornatomyia sp. (Davis 1990). However, it 

does little to control P. conjugaturn in the countries where it already occurs, so it does not 

appear to be a promising species to introduce elsewhere. 

Perhaps more valuable is the cecidomyiid fly Cleitodiplosis grarninis, a gall forming 

fly described from Brazil. The gall consists of the terminal leaves becoming clustered 

into an ovoid 30 x 20mm mass as a result of the upper internodes being greatly reduced. 

Thirty to forty sulphur-coloured larvae may be found in a single gall, in which they 

pupate. They appear in August and September (Barnes 1956). Barnes (1954b) comment- 

ed that no gall midge had, at that time, been recorded from the inflorescence of 

Paspalurn, although Parallelodiplosis paspali (from Java and India) and Lasioptera sp. 

had been recorded from the stems. 

The chrysomelid beetle Colaspis (= Maecolaspis) aerea also damages cocoa in 

Brazil (Fenonatto 1986) and hence would not be acceptable for introduction elsewhere. 

The seeds of P. conjugaturn are harvested by ants in some areas, but the impact of 

this on weed density in the field has not been established. Under experimental field con- 

ditions in Mexico, the pestiferous fire ant, Solenopsis gerninata, reduced P. conjugaturn 

seed densities by 97% or more but had no effect on the seed densities of Bidens pilosa 

(Carroll and Risch 1984). 

Comment 

As with Panicurn repens, so little is known about the natural enemies of Paspalurn conju- 

gatum that it is not possible to evaluate the prospects for its biological control. Again, 

there is the limitation that some closely related species in the genus Paspalum are of eco- 

nomic value for fodder (e.g. P. dilataturn (paspalum), P. distichurn (saltwater couch), P. 

plicatulum, P. scrobiculaturn (scrobic), P. vaginaturn (saltwater couch)). However, a fun- 

gus attacks the seeds of the first two species, producing the toxin, ergot, and most of the 

species are considered weedy in at least some situations, so there may be occasions 

where a conflict of interests would have to be resolved, should effective agents be dis- 

covered. The prospects for finding these are not perhaps very promising, since Holm et 

al. (1977) record P. conjugaturn as a weed in a number of countries in and around the 

Caribbean.


Table 4.21.1 Natural enemies of Paspalum conjugatum. 

Species Country References 

INSECTS 

Thysanoptera 

Haplothrips gowdeyi Hawaii Sakimura 1937 

Haplothrips paumalui Hawaii Sakimura 1937 

Coleoptera 

CHRYSOMELIDAE 

Colaspis (= Maecolaspis) aerea Brazil 

Diptera 

CEClDOMYllDAE 

Cleitodiplosis graminis Brazil 

Lepidoptera 

ARCTllDAE 

Creatonotos (= Amsacta) gangis 

HESPERllDAE 

Taractrocera ina 

PSYCHIDAE 

Brachycyttarus griseus 

FUNGI 

PYRALIDAE 


Cnaphalocrocis (= Marasmia) 

patnalis 


(= Nymphula depunctalis) 

Exserohilum paspali 

Leptosphaeria proteispora 

Myriogenospora atramentosa 

Physarum cinereum 

Sorosporium paspali 

BACTERIA 

Xanthomonas albilineans 

Ferronatto 1986 

Barnes 1956 

Philippines Catindig et al. 1993 

Australia Common & Waterhouse 198 1 

Guam, Hawaii, Malaysia, Davis 1990 

Vietnam, Philippines 

Philippines Abenes & Khan 1990 

Philippines Abenes & Khan 1990 

Philippines Bandong & Litsinger 1984 

Brazil Muchovej & Nesio 1987 

Hawaii Stevens 1925 

Brazil, USA, Venezuela Hanlin & Tortolero 1990 

Brazil Muchovej & Muchovej 1987 

Hawaii Stevens 1925 

Australia Persley 1973 

NEMATODES 

Rotylenchulus reniformis Trinidad Singh 1974 

VIRUSES 

sugarcane mosaic Hawaii, Japan, Taiwan Chen et al. 1989b, 

Holm et al. 1937, 

Ohtsu & Gomi 1985



(after Soe jani et a/. 1987)

Map 4.22 Passiflora foetida 

4.22 Passiflora foetida 


No searches have been made for natural enemies of P. foetida in its area of origin in 

South America, where it is not a weed. It is known to be attacked there by the larvae of 

some nymphalid (heliconiine) butterflies. 

However studies of the related P. tripartita, a serious forest weed in Hawaii, 

recorded upwards of 200 species of insects. It may thus be inferred that a similar study 

would reveal many insects attacking P. foetida. However, until a relevant study is carried 

out, it is not possible to evaluate the prospects for its successful biological control. 

I


4.22 PassifZora foetida L. 

Passifloraceae 

stinking passionflower, wild passionfruit; love-in-a-mist; ka thok rok (Thailand), 

timun padang (Malaysia), buah tikus (Indonesia) 

Rating 

+++ Msia 

11 ++ Brun 

+ Myan, Thai, Laos, Viet, Sing, Phil 

Indo 

Origin 

South America. Natural populations have been observed in the coastal mountain ranges 

in the State of Parana, Brazil (G.P. Markin pers. comm. 1993). 

Distribution 

Widespread throughout the tropics and serious in Southeast Asia; also a weed in the 

Pacific Region, West Africa and Central America. Introduced to Java a long time ago. 


A foetid, woody, annual or perennial vine, 1.5 to 6 m long; stem, cylindrical, densely 

hairy; tendrils arise next to leaves on the shaded side of the stem; leaves heart-shaped to 

three lobed, alternate, arranged helically, with long-stalked glands and long fine hairs on 

margins, producing a disagreeable smell when crushed; flowers white to lilac, bisexual. It 

flowers all year round, opening in the morning and closing before noon. The green to 

orange or red fruits are enclosed in lacy bracts. A large number of varieties occur 

(Wagner et al. 1990). 

Importance 

I! foetida is a weed of upland rice and other field crops. It occurs in wet areas or those 

where there is a pronounced wet season. It is common in plantations, rough pastures, 

roadsides and wasteland. 

In the Philippines it is sometimes used as a soil cover in coconut plantations to con- 

trol Imperata cylindrica grass or erosion. In Papua New Guinea it is planted between 

sweet potatoes to suppress Imperata. Young leaves are used in Surinam and Java as a 

vegetable. Seeds are flat, black, woody and enclosed in a sweet aromatic pulp (Swarbrick 

1981). Young fruit are cyanogenic. Stems and leaves are suspected of poisoning live- 

stock. 

P. foetida contains alkaloids and at least 10 flavonoids. One of the latter, ermanin, is 

a feeding deterrent to larvae of the nymphalid butterfly Dione juno which, in Colombia, 

do not attack P. foetida leaves, but eat large amounts of other PassiJlora species.

4.22 Passifora foetida 187 

The Passifloraceae contain about 12 genera and 600 species, most of which are ten- 

dril climbing vines native to warm regions of the world. The genus PassiJlora contains 

some 500 tropical and subtropical species, mostly from Central and South America. 

Several have edible fruits and attractive flowers, about 40 species have been cultivated, 

but fewer than 6 are fruit crops in the neotropics and only one, P. edulis (and its varieties, 

such as the yellow favicarpa), is economically important (Waage et al. 1981). P. ligu- 

laris is also cultivated in Malaysia (Ong and Ting 1973). A few species, such as P. foeti- 

da and P. lonchocarpa, are extremely foul smelling (Benson et al. 1976). Eleven species, 

including P. foetida and P. tripartita (= P. mollissima) (in Hawaii) are recorded as weeds 

in different parts of the world (Swarbrick 1981). Both P. foetida and P. tripartita are 

closely related taxonomically, whereas, P. edulis belongs to a different subgenus (Waage 

et al. 1981) and is the only economic crop at risk from oligophagous insects attacking 

P. foetida. 


Upwards of 200 insects are recorded attacking Passifloraceae in Central and South 

America. The most notable are heliconiine butterflies of the family Nymphalidae. Their 

larvae develop only on plants of the family Passifloraceae, with the single exception of 

Eueides procula, which will develop on the Turneraceae (Pemberton 1983, Waage et al. 

1981). only 5 of the 65 or so species of heliconiines are recorded as pests of Passiflora 

edulis, namely Agraulis vanillae, Dione juno, Dryas julia, Eueides aliphera and E. 

isabella, although larvae of a few other species are occasionally found on it (Waage et al. 

1981). It is apparent that heliconiine butterflies are well worth investigating for species of 

adequate host specificity to P. foetida. 

Little is known about the natural enemies of P. foetida (Table 4.22.1) and no 

attempts have been made at biological control. The passion vine butterfly Agraulis vanil- 

lae, an accidental introduction to Hawaii before 1977, is now widespread there. In addi- 

tion to attacking Passifora edulis, its larvae feed on the leaves of P. foetida, P. manicata 

and P. suberosa, but they seldom attack banana poka, P. tripartita, which is a serious for- 

est weed in Hawaii. P. foetida is widely distributed on Hawaii from sea level up to about 

500 m and a rainfall from 750 to 3000 mm. It occurs generally in highly disturbed areas, 

where it is a very minor component among other introduced species. On the west side, 

the taxon has red fruit and, on the east, green. It has very few natural enemies, with the 

exception of Agraulis vanillae. A. vanillae larvae are common but usually in small num- 

bers, although occasionally there are outbreaks that completely defoliate the plants (G.P. 

Markin pers. comm. 1993). A. vanillae is native to the Americas and ranges from 

Argentina up through Mexico to Florida, the Gulf States and California (Beardsley 1980, 

Bianchi 1982, 1983, Klots (1951)). In Hawaii it is attacked by a nuclear polyhedral virus 

which limits its numbers (G.P. Markin pers. comm. 1993), in California by Phorocera 

claripennis (Tachinidae) and in eastern USA by Brachymeria ovata (Nakahara 1977). 

The other species of heliconiine recorded as attacking P. foetida is Heliconius hecale, 

which is widespread in Central and South America and attacks a large number of


Passifloraceae (Benson et al. 1976, Waage et al. 1981). On the other hand, larvae of 

H. charithonia, H. cydno and H. erato did not develop on R foetida (Waage et al. 1981). 

In the Ivory Coast larvae of the pterophorid moth Sphenarches anisodactylus eat the 

leaves of R foetida, Lagenaria siceraria and Brillantaisia lamium. Although the moth 

also occurs in India and Japan it is not known from P. foetida there, but attacks two eco- 

nomic plants, the legumes lablab bean, Lablab purpureus and pigeon pea, Cajanus cajan 

(Bigot and Vuattoux 1979). Thus there is some uncertainty whether the host specificity of 

the African taxon is the same as that in Asia. 

The National Parks and Forest Service authorities in Hawaii have been carrying out 

searches for some years in South America for natural enemies of banana poka, R triparti- 

ta. Two insects from Colombia have been introduced to Hawaii (Gardner et al. 1992). 

One of these was the moth Cyanotricha necyria (Dioptidae), which was released in 1988, 

but did not become established (Casaiias-Arango et al. 1990, Julien 1992, Markin and 

Nagata 1989, Markin et al. 1989). In host specificity tests C. necyria did not oviposit on 

f? foetida, but the larvae could develop on its foliage (Markin and Nagata 1989). The 

fungus Colletotrichium gloeosporioides f. sp. clidemiae has been mass produced for lib- 

eration (E.E. Trujillo memorandum 1989). 

In Hawaii the fungus Fusarium oxysporum f. sp. passiflorue attacks P. foetida, 

P. tripartita and P. ligularis, but not R suberosa or the cultivated P. edulis f. jlavicarpa 

(Gardner 1 989). 

Table 4.22.1 Natural enemies of Passiflora foetida. 


INSECTS 

Hemiptera 

APHlDlDAE 

Aphis fabae Kenya polyphagous Bakker 1974 

Aphis gossypii Ivory Coast polyphagous De Wijs 1974 

Aphis spiraecola Ivory Coast De Wijs 1974 

Myzus persicae Japan polyphagous Yonaha et al. 1979 

Uroleucon compositae Kenya Bakker 1974 

Diptera 

AGROMYZIDAE 

Melanagromyza 

PO~YP~Y~~ 

Tropicomyia theae 

Lepidoptera 

NOCTUIDAE 

Helicoverpa zea 

(= H. obsoleta) 

. Heliothis virescens 

NYMPHALIDAE 

Agraulis vanillae 

f 

Australia polyphagous, including Kleinschmidt 1960, 1970 

Passijlora spp. 

Papua New polyphagous Spencer I990 

Guinea 

Sumatra polyphagous Den Doop 19 18 

Venezuela polyphagous Venturi 1960 

Central restricted to some Anon 1977, Beardsley 1980, 

America, Passijlora spp. Bianchi 1982, 1983, 

Hawaii Waage et al. 198 1 


FUNGI 

4.22 Passifloi-a foetida 189 


Heliconius hecale 

PTEROPHORIDAE 

Sphenarches 

anisodactylus 

Alternaria passiflorae 

Alternaria tenuis 

Colletotrichium 

gloeosporioides 

Fusarium oxysporum 

f.sp. passiflorae 

Haplosporella 

pass$oridia 

Hemphyllium sp. 

NEMATODE 


VIRUS 


passionfruit chlorotic 

spot 

passionfruit moiaic 

passionfruit ringspot 

potyvirus 

passionfruit woodiness 

potyvirus 

widespread in restricted to some 

Central & South Pass$ora spp. 

America 

Ivory Coast see discussion 

Hawaii 

Hawaii 

India 

Hawaii 

India 

Hawaii 

Australia 

Japan 

Papua New 

Guinea 

Hawaii, 

Malaysia 

Ivory Coast 

Australia, 

Kenya 

Waage et al. 198 1 

Bigot & Vuattoux 1979 

Raabe 1965 

Raabe 1965 

Mallikarjunaiah & Rao 1972 

Gardner 1989 

Pande 1980 

Raabe 1965 

Sauer & Alexander 1979 

Yonaha et al. 1979 

Van Velsen 196 1 

Ong & Ting 1973, 

Raabe 1965 

Brunt et al. 1990, 

De Wijs 1974 

Anon 1976, Bakker 1974, 

Brunt et al. 1990, 

Leggat & Teakle 1975




Map 4.23 Pennisetum polystachion 

4.23 Pennisetum polystachion 191 


This erect, tufted, non-stoloniferous grass, originated in Tropical Africa from where it 

has spread throughout Asia and Southeast Asia to the Pacific. 

Almost nothing is known of the natural enemies of P. polystachion or closely 

related species. It is not possible to evaluate the prospects for its biological control with- 

out a search for natural enemies in its region of origin.


4.23 Pennisetum polystachion (L.) Schultes 

(= Pennisetum setosum) 

Poaceae 

mission grass; feather Pennisetum; yaa khaehyon chop (Thailand), rumput 

gajah, rumput berus, rumput kuning, ekor kucing (Malaysia) rumput jurig 

(Indonesia) 

There are differences in opinion over the spelling of the specific name, polystachion or 

polystachyon, with the former being used here. In Africa, there are three subspecies 

P. polystachion polystachion, P. p. setosum (sometimes regarded as a true species) and 

P. p. atrichum. There is some evidence of crossing between the varieties of P. polysta- 

chion and the related Pennisetum hordeoides and the production of populations with dif- 

ferent chromosome numbers (Brunken 1979). 

Rating 

+++ Msia 

11 ++ Indo, Phil 

+ Thai, Laos, Camb, Viet 

Origin 

Tropical Africa. 

Distribution 

P. polystachion is widespread in the tropics of Africa and Asia, but also occurs in north- 

ern Australia and the Pacific. It rarely extends beyond 23"N or 23"s. In Africa it occurs 

mainly in the savanna and open areas in the forest zone of West Africa from Senegal to 

Cameroun and then south and east to Mozambique and Kenya (Brunken 1979, Kativu 

and Mithen 1988). 


P. polystachion is an erect, tufted annual or perennial grass, with fibrous roots, but no 

stolons. Its leaves are narrow and 5 to 45 cm long. Its flowering stems are sometimes 

branched, 50 to 300 cm tall, ending in a cylindrical yellow-brown flowering spike, 5 to 

25 cm long, bearing densely hairy, unequal bristles of two lengths, the longer 2 to 5 cm 

and the shorter lcm. 

Importance 

P. polystachion grows on dry lateritic soils and is often present along roadsides, in waste- 

lands and in upland crops. Propagation is by seeds, but regrowth can occur from dormant 

buds located at the base of the stems and from aerial nodes. 

It becomes dominant in upland tropical hillsides and croplands after forests have

4.23 Pennisetum polystachion 193 

been cleared, or when shifting cultivation or subsistence agriculture have been practised 

(Holm et al. 1977). Because of the rapid germination of its wind-dispersed seeds and its 

aggressive and highly competitive growth, it rapidly takes over wastelands. Since a sin- 

gle cultivation rarely kills enough of the weed to provide control, it often impedes further 

use of areas for crops. 

In Indonesia it was first observed in 1972 (Titrosoedirdjo 1990). It is now an impor- 

tant weed of rubber and occasionally a problem in upland rice. 

P. polystachion is thought to have reached Malaysia via Thailand as recently as the 

early 1980s, infesting at least 10 km2 of roadsides and is now widely distributed, occur- 

ring up to an altitude of 900 m (Baki et al. 1990). It is now a major weed in rubber, oil 

palm, sugarcane, orchards, vegetables and upland rice (Titrosoedirdjo 1990). 

In the Philippines P. polystachion is able to compete effectively even with blady 

grass Imperata cylindrica in grassy fields in Central Luzon and in rubber plantations in 

west Java (Titrosoedirdjo 1990). 

P. polystachion is a good fodder grass when young and makes excellent hay. 


Very few natural enemies of P. polystachion appear to have been recorded in the litera- 

ture (Table 4.23.1). The only species of possible relevance are several gall midges from 

Africa, but very little is recorded of their biology. Three species of gall midge have been 

reared from P. polystachion in the Gold Coast. One is similar to the pestiferous sorghum 

midge Contarinia sorghicola (but may be different), the second belongs to the Trifila 

group and the third belongs to the Lasiopterariae (Barnes 1954a,b, Geering 1953). Three 

species of gall midge have been described from ears of Pennisetum in southern India, 

Cecidomyia penniseti (from P. glaucum = P. typhoideum), Geromyia (= Itonida) penniseti 

(from P. cenchroides) and Geromyia (= Itonida) seminis and an unidentified species from 

the stems. Of these, G. penniseti may be predaceous (Barnes 1954b, Felt 1920, 1921). 

In Madagascar there is a gall midge (?Cecidomyia sp.) whose larvae live in the 

inflorescence of Pennisetum (no species given) and in Sudan a gall midge, possibly 

Geromyia seminis, has been reared from the ears of Pennisetum (no species given) 

(Barnes 1954b). 

The larvae of the Brazilian satyrid butterfly Eryphanis polyxena were bred in the 

laboratory on P. polystachion (= P. setosum) (Dias 1979), but damage is not reported 

from the field. 

Comment 

There is little doubt that a range of natural enemies attacking f? polystachion would be 

found if a search were made in Tropical Africa. 

Several other weedy species of Pennisetum also originated in Africa, in addition to 

three or more species that have at least some desirable attributes. Perhaps the best known is 

kikuyu grass, P. clandestinum, which is a valuable fodder during the warmer months, 

although it is a weed in some situations and its nitrate levels can be toxic to grazing animals. 

?? glaucum (= P. americanum), pearl millet, is used as food in some areas. ?? purpureum,

194 


elephant or napier grass, can grow to 3 m, is valuable for fodder when young and can be 

used as fuel when old. However, it can also be a weed, as in rubber in Malaysia. Little 

has been recorded of the natural enemies attacking these species of Pennisetum. The 

most interesting are three cecidomyiid gall midges in India, Geromyia penniseti, G. sem- 

inis and Mycodiplosis indica from Pennisetum glaucum (= P. typhoideum) and Geromyia 

seminis from Pennisetum cenchroides (Barnes 193 1, Felt 1920). 

Table 4.23.1 Natural enemies of Pennisetum polystachion. 

Species 

INSECTS 

Diptera 

CEClDOMYllDAE 

Cecidomyia penniseti 

Contarinia ?sorghicola 

Contarinia sp. 1 




Geromyia (= Itonida) penniseti 

Geromyia (= Itonida) seminis 

Lepidoptera 

HESPERllDAE 

Parnara bada bada 

NYMPHALIDAE 

Eryphanis polyxena 

PYRALlDAE 

Cnaphalocrosis medinalis 

FUNGI 

Bipolaris papendor-i 

Gloeocercospora sp. 

Helminthosporiurn rostratum 

Phakospora apoda 

Puccinia chaetochloae 

Puccinia substrata 

Pyricularia oryule 

Spacelotheca penniseti 

Country 

India 

Gold Coast 

Gold Coast 

Gold Coast 

Madagascar 

India 

India 

India. Sudan 

Malaysia 

Brazil 

Malaysia 

Brazil 

References 

Barnes 1954b 

Barnes 1954b 

Bames 1954b 

Barnes 1954b 

Barnes 1954b 

Bames 1954b 

Barnes 1954b 

Barnes 1954b 

C.L. Tan pers. comm. 1993 

Dias 1979 

C.L. Tan pers. comm. 1993 

H.C. Evans pers. comm. I992 

H.C. Evans pers. comm. 1992 

Thite & Chavan 1977 




Prabhu et al. 1992 

H.C. Evans pers. comm. I992




Map 4.24 Pistia stratiotes 

4.24 Pistia stratiotes 

Pista stratiotes 

Water lettuce is a widespread, floating water weed, which probably originated in South 

America. 

The host specific South American weevil, Neohydronomus aflnis, has been 

established readily in six countries and, in all, has produced substantial to excellent con- 

trol. The moth, Samea multiplicalis, which attacks P. stratiotes and Salvinia spp., has 

been established in Australia but its impact has not been evaluated. 

In Thailand, classical biological control has not been attempted, but mass rearing 

and release of the native noctuid moth Spodoptera pectinicornis has replaced the use of 

herbicides. 

The prospects are excellent for classical biological control of P. stratiotes in 

countries where it is still regarded as an important weed.


4.24 Pistia stratiotes L. 

Araceae 

water lettuce; chak thom (Cambodia), apoe apoe, apon apon (Indonesia), 

kiambang besar (Malaysia), chok, jawg (Thailand), beo cai (Vietnam) 

Rating 

++ Thai, Laos, Camb, Viet, Phil 

14 + Myan, Msia, Brun, Indo 

Sing 

Origin 

The origin of Pistia stratiotes is unknown, although the number of host specific insects 

present there (Table 4.24.1) suggests South America. However, there is also a host spe- 

cific noctuid moth that ranges from India to Papua New Guinea. Dray and Center (1992) 

examine the various theories concerning the area of origin of water lettuce. 

Distribution 

Water lettuce occurs very widely as a troublesome water weed between the tropics of 

Capricorn and Cancer, particularly in Africa, Asia, Southeast Asia and the Caribbean. 

Pliny refers to its use in Egypt in AD77 (DeLoach et al. 1979, Holm et al. 1977). It is 

absent from a number of Pacific countries although recorded as a weed in Papua New 

Guinea, Solomon Is, Guam, New Caledonia and Hawaii. 


Pistia stratiotes is a free-floating, perennial monocotyledon, with a tuft of fibrous feath- 

ery roots up to 1 m long. Numerous secondary roots may be up to 4 cm in length. Stolons 

up to 60 cm are produced from the base of the plant and develop into new plants. Leaves 

are pale green, upright, 2.5 to 15 cm long, broad at the top and tapered at the base. They 

are prominently veined beneath and form a rosette. They are spongy in texture and bear 

numerous fine, water-repelling hairs on both sides. The flowers are bisexual, inconspicu- 

ous, green, surrounded by tubular bracts and arise from the centre of the rosette. It is said 

that water lettuce does not flower in Thailand but a small number of flowers have been 

observed (B. Napompeth pers. comm. 1993). It flowers in the Philippines, Australia, 

Africa and USA. The fruit is berry-like and green and contains 4 to 12 small brown seeds 

which can float on the water for up to 2 days. There are as many as 9 varieties of water 

lettuce (Neal 1965). 

Importance 

The free-floating plants are found in reservoirs, ponds, in marshes along the edges of 

large tropical lakes and in slow-moving or stagnant waters. They multiply rapidly and

4.24 Pistia stratiotes 199 

can block streams, interfere with fisheries and hydroelectric generating plant and bank up 

at dams, bridges and culverts, leading to increased flooding problems. Despite earlier 

claims (Holm et al. 1977), unlike water hyacinth, water lettuce does not increase water 

loss through evapotranspiration (Lallana et al. 1987). Together with water hyacinth it is a 

common and important component of the dense aggregations of free-floating vegetation, 

known as sudds. It grows best at pH 4, whereas water hyacinth produces greatest dry 

weight at pH 7 (Holm et al. 1977). 

Water lettuce plants act as a substrate for sandfly larvae (Ceratopogonidae) and lar- 

vae and pupae of the disease-transmitting mosquito genus Mansonia obtain their oxygen 

by attaching to Pistia roots. P. stratiotes is an important weed of irrigated rice, floating 

into paddy crops, taking root in the soil and competing much like other weeds. 

On the other hand, it has been used as human food in India during famines and is 

still fed to pigs and ducks. It is said to have some medicinal value as a cure for skin dis- 

eases and dysentery, as a laxative, to treat asthma and, its ash rubbed into the scalp, as a 

treatment for ringworm. 


AUSTRALIA 

Although water lettuce was first recorded in the Northern Territory only in 1946 it was 

already an important weed in some locations in Queensland before the introduction of 

Neohydronomus afJinis, although plants in the Northern Territory were rarely thrifty. 

Heavy damage was observed there by larvae of the moth Parapoynx (= Nymphula) tene- 

bralis, which lays it eggs on the leaves. Newly emerged larvae excise a portion of leaf to 

make a protective case in which they shelter while feeding and moving around the plant. 

These larvae also attack Salvinia molesta, as do larvae of the related Parapoynx 

(= Nymphula) turbata (Gillett et al. 1988). In Thailand this species attacks water lettuce 

(Napompeth 1982), so it presumably has the same habit in the Northern Territory. A bug, 

Nisia nervosa (= N. atrovenosa) feeds on water lettuce as it does in India (Gillett et al. 

1988, Joy 1978). 

CENTRAL AND SOUTH AMERICA 

Bennett (1975) listed 12 species of phytophagous insects that had been reared from Pistia 

stratiotes and Cordo et al. (1981) added one more. Particularly notable (Table 4.24.1) is 

the group of South American weevils which are generally confined to water lettuce, 

although adults of several may produce minor attack on some of the nearby aquatic 

plants whose stems and leaves nevertheless would be too small to support larval develop- 

ment. Preliminary host range studies suggest that several may be specific enough to be 

employed for biological control, although adequate information is available only for 

Neohydronomus afinis. This has been successfully established in several countries (see 

later). Larvae of the small pyralid moth Samea multiplicalis, which occurs from 

Argentina to the southeastern United States, feed on the growing buds of water lettuce 

and sporadically cause very heavy damage and dieback of the plants (Cordo et al. 1978, 

1981, DeLoach et al. 1976, 1979).


Table 4.24.1 Natural enemies of Pistia stratiotes. 


INSECTS 

Hemiptera 

APHlDlDAE 

unnamed 

Rhopalosiphum nymphaeae 

COCCIDAE 

Planococcus citri 

LYGAEIDAE 

Valtissius sp. 

MEENOPLllDAE 

Nisia nervosa 

(= N. atrovenosa) 

Orthoptera 

ACRlDlDAE 

Paulinia acuminata 

TETRlGlDAE 

Criotettrix sp. 

Coleoptera 

CURCULIONIDAE 

Argentinorhynchus bennetti 

Argentinorhynchus breyeri 

Argentinorhynchus bruchi 

Argentinorhynchus minimus 

Argentinorhynchus squamosus 

Neohydronomus aflnis 

Neohydronomus elegans 

Neohydronomus pulchellus 

Ochetinu bruchi 

Pistiacola cretatus 

Pistiacola fasciatus 

Pistiacola sp. nr nigrirostris 

Lepidoptera 

ARCTllDAE 

Spilosoma virginica 

NOCTUIDAE 

Erastroides curvifascia 

Proxenus hennia 

,Spodoptera pectinicornis 

Thailand 

Florida 

Nigeria, Trinidad 

Argentina 

Australia. India 

South America 

Thailand 

Mexico, Venezuela 

Argentina, Paraguay 


Venezuela 


South America 

Honduras, Cuba 

Trinidad to Argentina, Cuba 

Argentina 

Argentina, Brazil 

Central and South America 

Argentina 

Florida 

India 

Indonesia 

India, Bangladesh, Thailand 

Napompeth 1990a 

Ballou et al. 1986, Bennett 

1975, Joy 1978 

Bennett 1975 

Bennett 1975 

Gillett et al. 1988, Joy 1978 

Bennett 1966, 1975 


O'Brien & Wibmer 1989a,b 


Cordo et al. 1978 


Cordo et al. 1978 

DeLoach et al. 1976 

O'Brien & Wibmer 1989c 

Bennett et al. 1975, 

O'Brien & Wibmer 1989c 

Cordo et al. 198 1 


Wibmer & O'Brien 1989 


Thompson and Habeck 1988 

Chaudhuri & Janaki Ram 

1975 

Mangoendihardjo & Nasroh 

1976 

Alam et al. 1980, George 

1963, Napompeth 1990a, 

Sankaran 1974, Sankaran & 

Rao 1972, Sankaran et al. 

1964 


MITES 

FUNGI 



PYRALlDAE 

Argyractis subornata 

Elophila responsalis 

Parapoynx (= Nymphula) 

diminutalis 


tenebralis 


turbata 

Petrophila drumalis 

Samea multiplicalis 

Synclita obliteralis 

Hydrozetes subornata 

Cercospora canescens 

Cercospora sp. 

Phyllosticta stratiotes 

Sclerotium rolfsii 

Brazil 

India, Indonesia 

Thailand 

Australia 

Australia, Thailand 

Florida 

southern USA, Trinidad, 

northern South America 

Florida 

Australia 

Australia 

India 

India 

India 

Fomo 1983 


Mangoendihardjo et al. 1977, 

Sankaran & Rao 1972, 

Subagyo1975 

Napompeth 1990a, 

Suasa-Ard 1976 

Gillett et al. 1988 

Gillett et al. 1988, 

Napompeth 1990a, 

Suasa-Ard 1976 

Dray et al. 1988 

Bennett 1966, 1975, 

Bennett et al. 1975, 





Bennett 1975, 

Nag Raj and Ponappa 1966 

Bennett 1975 

Bennett 1975 

UNITED STATES 

Dray et al. (1988) recorded larvae of three species of moth, a mealy bug, a leafhopper 

and an aphid on water lettuce which has been present in Florida for at least 200 years 

(Thompson and Habeck 1988). Only one of these insects, a root feeding moth, was con- 

sidered to be possibly host specific. This was later identified as the pyralid moth 

Petrophila drumalis: the two other moths were Samea multiplicalis and Synclita obliter- 

alis (Dray et al. 1989). The aphid was probably Rhopalosiphum nymphaeae, a well 

known transmitter of a number of economically important viruses. It has been recorded 

to cause dieback of water lettuce in Nigeria (Pettett and Pettett 1970). This aphid was 

reported to be widespread on water lettuce in Florida (Ballou et al. 1986). An aphid, pos- 

sibly the same species, transmitted a virus that caused widespread dieback of Z? stratiotes 

on Lake Volta in Ghana (Okali and Hall 1974), although serious dieback has not been 

reported in Florida. The non-specific larvae of the arctiid moth Spilosoma virginica was 

also common on water lettuce in Florida (Thompson and Habeck 1988).


INDIA 

Larvae of the moth Spodoptera pectinicornis cause extensive damage to Pistia. On aver- 

age, a single larva can consume the leaves of two Pistia plants during its developmental 

period of 15 to 20 days. Some 100 larvae developing from an average egg mass destroy 

all Pistia leaves within an area of 1 m2 and, during peak abundance in the field, the num- 

ber of larvae per m2 of Pistia surface was always higher than this (George 1963, 

Sankaran and Ramaseshiah 1974). The bug Nisia nervosa successfully completes its life 

cycle on Pistia, but is reported as a minor pest of rice (Joy 1978). 

INDONESIA 

In Java and Sulawesi, water lettuce is attacked by larvae of the noctuid moth Proxenus 

hennia which appears to be specific (Mangoendihardjo and Nasroh 1976). Other species 

found attacking it were Elophila (= Nymphula) responsalis, Spodoptera mauritia, an 

aphid and a cicadellid (Zygina sp.) (Mangoendihardjo and Syed 1974, Mangoendihardjo 

et al. 1976, 1977, Syed et al. 1977). 

THAILAND 

Water lettuce is attacked by several insects (Table 4.24.1), of which only the pygmy 

grasshopper Criotettrix sp. and the native water lettuce moth Spodoptera pectinicornis 

are capable of inflicting serious damage. In certain areas where the density of Criotettrix 

was as high as 100 per m2 considerable suppression of the weed occurred. Both adults 

and nymphs were able to walk on the surface of the water and were observed to attack 

also the water fern, Salvinia cucullata (Napompeth 1982). The extensive damage that can 

be caused by Spodoptera pectinicornis is discussed later. 


AUSTRALIA 

The first attempt to bring about classical biological control of Pistia stratiotes was the lib- 

eration of adults and larvae of Neohydronomus afinis in 1982 near Brisbane. Within two 

months of release, plants were rotting and sinking and, by eight months, about one third 

of the plants in a dam were chlorotic and some had been destroyed. Severely damaged 

plants produced short stolons terminating in small plantlets before sinking and dying, but 

these plantlets failed to grow to the size of their parents before, in turn, becoming severe- 

ly damaged, producing plantlets and then sinking. Continued weevil attack led initially to 

an increase in the number of plants, but a decrease in their size and dry weight. Before 

long, few water lettuce plants remained (Harley et al. 1984). The moth Samea multipli- 

calis was liberated in Australia in 1981, primarily against Salvinia molesta, on which it 

became established. However, within four years, its effectiveness was restricted by proto- 

zoan disease and three hymenopterous parasitoids (Thomas and Room 1986). It presum- 

ably attacks Pistia stratiotes also, although there seems to be only one observation of it 

doing so. This was at Townsville (D.P.A. Sands pers. comm. 1993). 


The moth Spodoptera pectinicornis attacks water lettuce, but is unable to prevent its 

increase when the plant is freed from competition by the biological control of Salvinia or

Table 4.24.2 Liberations for the biological control of Pistia stratiotes. 


Species Where From When Result References 

Coleoptera 

CURCULIONIDAE 

Neohydronornus a#nis Australia Brazil 1982 + Harley et al. 1984, 1990 

Botswana Brazil via 1988 + Chikwenhere & Forno 1991 

Australia LW. Fomo pers. comm. 1993 

Papua New Brazil via 1985 + Chikwenhere & Forno 199 1, 

Guinea Australia Harley et al. 1990, 

Laup 1987b 

South Africa Brazil via 1985 + Cilliers 1987, 1989b 

Australia 

United States Brazil via 1987 + Center et al. 1989, 

of America Australia Thompson & Habeck 1988, 


Zambia Zimbabwe about + P. Room pers. comm. 1993 

1990 

Zimbabwe Brazil via 1988 + Chikwenhere & Fomo 1991 

Australia 

Lepidoptera 

NOCTUIDAE 

Spodoptera pectinicornis Florida Thailand 1990 ? Center et al. 1989, 

Julien 1992 


PYRALl DAE 

Sarnea rnultiplicalis Australia Brazil 198 1 + Forno 1987, Room et al. 1984 

Eichhornia. Neohydronomus afinis was successfully established in the Sepik River sys- 

tem in 1985, but its impact is yet to be recorded (Laup 1987b). 

THAILAND 

Although no introductions of biological control agents for Pistia stratiotes have been 

made in Thailand, the mass rearing and release of the native noctuid moth Spodoptera 

pectinicornis has replaced the use of herbicides for this weed. Under laboratory condi- 

tions mixed instar larvae at the rate of 300 or more per m2 gave as fast and effective con- 

trol as any herbicide. In the field a substantial initial release of larvae, followed by one or 

two additional releases at two-week intervals has led to complete control within 6 to 10 

weeks. Thus, a 4.5 km2 infestation of water lettuce was controlled in 6 weeks at Sri 

Nakarint Dam in 1978 and a 10 km2 infestation in 1982 (Napompeth 1982). S. pectini- 

cornis occurred throughout the year and in all infestations of Pistia (Suasa-Ard and 

Napompeth 1982). 

UNITED STATES 

Neohydronomus afinis was released in Florida in 1987, became established readily, mul- 

tiplied rapidly and soon spread from the release sites to cause considerable damage to 

water lettuce (Dray et al. 1990). In one release site the Pistia population was reduced 

from 50 to less than 5 acres in 2 years and, in another, a 10 acre site was virtually cleared


in 3 years. However, in a third site, little effect was noted. It was postulated that this 

might be due to the presence of a different genetic strain of P. stratiotes, which had a far 

greater seed production than that at the other two sites (Dray and Center 1992). 

Spodoptera pectinicornis has also been established in Florida (Center et al. 1989; 

Napompeth 1 990a). 

BOTSWANA 

N. afinis was released on the Linyanti R at the Selinda spillway in 1988. Excellent con- 

trol was achieved within 12 months (I.W. Forno pers. comm. 1993). 

SOUTH AFRICA 

Neohydronomus afinis was released in Kruger National Park and a water lettuce infesta- 

tion in a motionless water body was completely controlled within 10 months (Cilliers 

1987, 1989a,b). The weevil has been less successful on fast-flowing rivers where plants 

infested with weevil larvae are continually washed down stream and replaced by unin- 

fested plants from higher up. However, even under these circumstances, up to 90% of 

plants showed signs of feeding damage (Cilliers 1991 b). 

ZAMBIA 

N. afinis was already established by natural spread at Kafubu Lake when N. afinis from 

Zimbabwe was liberated about 1990 and by 1992 there were only scattered plants of 

Pistia but no mats (P.M. Room pers. comm. 1993). 

ZIMBABWE 

Neohydronomus afinis was released in 1988, was well established in 4 months and, with- 

in a year, water lettuce was no longer a problem in the Manyame River (Chikwenhere 

and Forno 1991). 

Major natural enemies 

Argentinorhynchus bmchi Coleoptera: Curculionidae 

This yellow spotted weevil (4.7 mm long) is known from Argentina, Bolivia, Paraguay 

and Uruguay (O'Brien and Wibmer 1989a,b). Although it is rare, it has the potential to 

cause heavy damage to water lettuce. Under laboratory conditions adults ate 1 cm2 of 

leaf surface per day, producing some 10 oval holes all the way through the leaf. Adults 

feed mostly by night and generally on medium-aged leaves. Field collected females laid 

on average 1575 eggs among the dense hairs on the leaf surface. Eggs hatch in 7.6 days. 

First instar larvae enter the leaf and feed on the spongy leaf tissue and second and third 

instars in the crown. Fourth instar larvae could not be reared: in the laboratory they left 

the plant and drowned. Adults fed and oviposited only on water lettuce and, except for 

slight feeding on Spirodela, of the 26 plant species tested, larvae only developed on water 

lettuce. In the laboratory 6 larvae per plant killed water lettuce within a month. It was 

suggested that egg predation may account for the rarity of A. bruchi; also that the diffi- 

culty experienced in rearing the fourth instar larvae may indicate that special conditions 

are required, lack of which may reduce survival (Cordo et al. 1978).


Neohydronomus aflnis Coleoptera: Curculionidae 

This mottled, brown-grey weevil was earlier confused in the literature with the closely 

related N. pulchellus. It occurs naturally in Argentina, Brazil, Colombia, Paraguay, Peru, 

Uruguay and Venezuela (O'Brien and Wibmer 1989~). Adults (males 1.8 mm, females 

2.1 mm long) feed on the leaves of Pistia stratiotes and mine the tissues: they do not 

appear to attack the crown or roots. Females lay about 1 egg per day beneath the leaf epi- 

dermis, usually on the upper surface near the margin. Eggs hatch after 3 to 4 days and 

larvae tunnel through the leaf tissues to complete development in 1 1 to 14 days. Pupation 

occurs in small pockets in the leaf tissues and adults emerge after about 4 days. The peri- 

od from egg to adult varies from 4 to 6 weeks, but there are only 3 generations a year in 

Argentina (December, February to March and June). Overwintering probably occurs in 

the adult stage (DeLoach et al. 1976). 

N. afJinis is very destructive under laboratory conditions. Maximum damage 

occurred in midsummer in Argentina, when peak populations of 200 to 600 per m2 pro- 

duced 1.6 feeding spots per cm2 of leaf surface. Adult N. afinis are occasionally para- 

sitised by nematodes in Argentina (DeLoach et al. 1976). 

N. affinis is highly specific to water lettuce, as shown by tests in Zimbabwe 

(Chikwenhere and Forno 1991), South Africa (Cilliers 1989b), Florida (DeLoach et al. 

1976, Dray et al. 1990, Thompson and Habeck 1988, 1989) and Australia (Harley et al. 

1990); and also by absence of reported damage to economic plants in any of the countries 

to which it has been introduced. 

Pistiacola cretatus Coleoptera: Curculionidae 

This brown 2.3mm long weevil, earlier known as Onychylis cretatus, occurs in Argentina 

and Brazil (Wibmer and O'Brien 1989). Adults feed mainly on the upper surface of the 

leaves of P. stratiotes and oviposit into the leaf tissue. The slender larvae tunnel into the 

denser tissues of the basal third of the leaf and also into the crown. Pupation occurs with- 

in the spongy part of the leaf. In the field, adult P. cretatus were found only on water let- 

tuce (Cordo et al. 1981). 

Samea multiplicalis Lepidoptera: Pyralidae 

This brown moth with dark markings and a wingspan of about 17 mm occurs from 

Florida to Argentina. 

Up to 290 eggs are laid per female, mainly on the upper surface of the leaves. These 

hatch after 4 days and the larvae construct a silken canopy under which they feed, or they 

may tunnel into the leaves to feed on the spongy tissues: they also eat the buds. After 5 or 

6 instars in the course of 16 days, they pupate in silken cocoons, to emerge as adults 5 

days later. Adults live up to 7 days (DeLoach et al. 1979, Knopf and Habeck 1976, Sands 

and Kassulke 1984). S. multiplicalis has three main hosts in Florida, Pistia stratiotes, 

Azolla caroliniana and Salvinia rotundifolia and it may occasionally attack Eichhornia 

crassipes. Oviposition is highest on P. stratiotes. Although medium to large larvae fed on 

a number of plants under laboratory conditions, S. multiplicalis has never been reported


as a pest of cultivated plants in Argentina or Brazil (DeLoach et al. 1979). It passed strict 

host specificity tests in Australia, larvae completing development on l? stratiotes, Azolla 

pinnata and Salvinia molesta. Larvae that had fed first on S. molesta were unable to com- 

plete their development on water lettuce, although they produced minor leaf scars. S. 

multiplicalis was released in Australia, but primarily against Salvinia molesta (Sands and 

Kassulke 1984). 

Samea multiplicalis has 3 generations a year in the field in Argentina, with popula- 

tion peaks in December, February and May, when populations reach a maximum of 5 lar- 

vae per plant. In laboratory tests females laid 99.3% of their eggs on P. stratiotes. Larvae 

caused heavy, but sporadic, damage to water lettuce in the field. However, in most years, 

populations were held at low levels, apparently by parasitoids (Apanteles sp. and 

Podogaster sp.) (DeLoach et al. 1979). In Florida 52% parasitisation was recorded, 

42.7% by three species of Hymenoptera (Agathis sp., Apsilops sp. and Temelucha ferrug- 

inae) and 9.3% by a tachinid fly (Lixophaga sp.) (Knopf and Habeck 1976). Nosema sp. 

was detected in Australia in some larvae from Brazil and the culture was freed of these 

before release (Sands and Kassulke 1984). 

Spodoptera pectinicornis Lepidoptera: Noctuidae 

This moth ranges over an extensive area from India through Sri Lanka, Thailand and 

Indonesia to Papua New Guinea. Eggs are laid in masses of 70 to 120 on the undersur- 

face of the Pistia leaf near its edge. They hatch in 40 to 60 hours to produce pale green 

larvae that burrow in the leaf parallel to the longitudinal veins. After some 20 days the 

1.5 to 2 cm long larvae pupate, to emerge two to three days later as small silvery brown 

moths about lcm long (George 1963). In Thailand the period from egg to adult averaged 

30 days and females laid an average of 666 eggs. Host specificity tests showed that it 

would develop only on Pistia stratiotes (Suasa-Ard 1976). Napompeth (1990a) reports 

that it is relatively simple to mass rear in the laboratory and to distribute in the field. 

Details are available of its rearing and ecology in Thailand (Suasa-Ard 1976, Suasa-Ard 

and Napompeth 1978). It was mass reared and released in Florida after tests showed that 

it was sufficiently host specific (Center et al. 1989, Napompeth 1990a). 

Comment 

Pistia stratiotes is seldom more than a minor component of the floating weed mass when 

either Eichhornia crassipes or Salvinia molesta or both are present. However, when it 

occurs alone or when the strong competition from these two weeds is greatly reduced by 

their effective biological control, water lettuce can increase rapidly to occupy the vacated 

water surface. Since damaging biological control agents are available for all three weeds, 

it is sensible to embark on a biological control program for them all, either at the same 

time or in sequence. 

Adequate biological control of water lettuce has been achieved by the introduction 

of the weevil Neohydronomus alone. However, if an even greater degree of control is 

desired, there are, in South America, additional species of weevil and also several moths


that appear to be well worthwhile investigating further. This is particularly so, as some 

are known to be heavily attacked by natural enemies and, if introduced without these, 

would be expected to be even more effective. 

It can be concluded with some confidence that water lettuce is a promising candi- 

date for biological control.




Map 4.25 Portulaca oleracea 

4.25 Portulaca oleracea 209 

Portulaca oleracea is a serious weed throughout tropical, subtropical and temperate 

areas, attaining high overall pest status more because of its very widespread importance 

than by being amongst the top few weeds in any one country. 

About 14 of the 140 or so species of insects that are known to attack it appear to 

be restricted to the genus Portulaca and probably several to P. oleracea or its very close 

relatives. In their native ranges 4 leaf-mining or gall-forming flies, 1 leaf-mining moth, 1 

leaf-mining sawfly and 2 weevils all cause considerable damage and show high specifici- 

ty to P. oleracea. 

If this group of phytophagous insects is not already present, the establishment of 

several without their own natural enemies should lead to a significant lowering in the 

weed status of I? oleracea. 

Portulaca oleracea is an attractive target for an attempt at biological control.


4.25 Portulaca oleracea L. 

Portulacaceae 

pigweed, purslane; gelang, krokot (Indonesia); gelang pasir, segan (Malaysia); 

phak bia yai (Thailand), mya byit, mye byet (Myanmar); kbet choun (Cambodia); 

golasiman, ulasiman (Philippines), rau Sam (Vietnam) 

Rating 

+++ Phil 

10 ++ Thai 

+ Myan, Viet, Msia, Sing, Indo 

Much of the material in this dossier is summarised from the account in Waterhouse 

(1993b). 

Origin 

Uncertain; possibly Central America, but see comment. 

Distribution 

Very widespread in tropical, subtropical and temperate regions of the world, including 

Southeast Asia, Australia and the Pacific. 


Pigweed is a fleshy annual herb, reproducing by seed, or by stem-fragments rooting 

when lying on moist soil. The stems are succulent, often reddish and 0.2 to 0.5 m in 

length. The stems and leaves are smooth and fleshy and form mats. In sunlight the plants 

are prostrate, but in partly shaded situations they may attain a height of 0.5 m. The leaves 

are alternate, flowers are yellow, sessile, self-pollinated and either occur singly, or sever- 

al may occur together, in the leaf clusters at the ends of branches. They open on sunny 

mornings and later produce numerous tiny (0.5 mm diameter) black seeds. 

Importance 

P. oleracea was ranked 9th of the world's worst weeds, being recorded in 45 crops in 81 

countries (Holm et al. 1977). With a rating of 10 in Southeast Asia, it ranked equal 32nd 

in the region; also 6th in the Pacific and 49th in Australia (Waterhouse 1993a,b). In 

Southeast Asia, it is particularly important in many upland crops, including vegetables, 

rice, maize, sorghum, groundnuts and sugarcane. It is drought hardy, colonising waste 

places and bare areas but thrives in moist fertile soils. There are many ecological types, 

some of which are occasionally used as a vegetable. In the Philippines up to 10000 and 

in Norih America up to 243 000 seeds are produced per plant. The tiny seeds are spread 

by wind, water, as a contaminant of the seeds of crops and by birds, surviving passage 

through the digestive tract. They also survive burial for long periods and germinate best 

above 30°C, but poorly below 24°C.


Pigweed does not compete well with other weeds. It is successful because it estab- 

lishes rapidly after soil disturbance and may flower and seed before being outcompeted 

by taller plants. The succulent leaves and stems are rich in oxalates and nitrates and have 

been implicated in livestock deaths. The succulent leaves of some strains have been used 

as human food (Miyanishi and Cavers 1980). 


The 138 insect species that have been recorded attacking P. oleracea were listed by 

Waterhouse (1993b), most of them from Central and South America (Bennett and 

Cruttwell 1972) and USA (Romm 1937). A few additional records are now listed in table 

4.25.1. Most of these species are known to be (or suspected of being) polyphagous and 

many are pests. Nevertheless, table 4.25.2 lists 14 insects that, so far as known, are 

restricted to P. oleracea, or at least to the genus Portulaca. Eight of these appear to have 

originated in the Americas, 2 each in Africa and India, and 1 each in France and 

Southeast Asia. With the exception of the weevil Ceutorhynchus portulacae, described 

from P. oleracea in Java, there do not appear to be any reports of native insects which 

might possibly be restricted to pigweed in Southeast Asia, the Pacific or Australia. 

Table 4.25.1 Natural enemies of Portrclaca oleracea: additional insect records to those 

of Waterhouse (1 993b). 

Species Country Other hosts References 

Hemiptera 

APHlDlDAE 

Myzus persicae 

ClCADELLlDAE 

Circulifer haematoceps 

Orosius orientalis 

(= 0. albicinctus) 

LYGAEIDAE 

Nysius cymoides 

Nysius vinitor 

Coleoptera 

CURCULIONIDAE 

Hypurus bertrandi 

Diptera 

AGROMYZIDAE 

Liriomyza caulophaga 

Liriomyza trifolii 

1 

Australia polyphagous author's record 

Israel polyphagous Klein & Raccah 1991 

India polyphagous Kooner & Deol 1982 

Italy jojoba Parenzan 1985 

Australia polyphagous Elshafie 1976, Ramesh & 

Laughlin 1984 

Australia, India, specific R.E. McFadyen pers. comm. 

Guam, Northern 1993,Zaka-ur-rab 199 1, 

Marianas Zimmerman 1957 

Australia Beta vulgaris R.E. McFadyen pers. comm. 

var. cicla 1993 

USA polyphagous Chandler & Chandler 1988 





Lepidoptera 

NOCTU IDAE 

Agrotis ipsilon India polyphagous Das & Ram 1988 

Neogalea (= Spodoptera) 

sunia Nicaragua polyphagous Savoie 1988 

Spodoprera eridania Nicaragua polyphagous Savoie 1988 

Spodoptera exigua Nicaragua polyphagous Savoie 1988 

PY RALl DAE 

Loxostege bijdalis N. America cotton Allyson 1976 

The host specificity has been investigated of 5 of the 6 species of Diptera, 

Lepidoptera and Hymenoptera listed in table 4.25.2, but, except for Baris arctithorax and 

Hypurus bertrandi, both of which appear to be adequately specific (see later), little is 

known about that of the 8 weevils. 

Several of the fungi listed in table 4.25.3 are reported to damage P. oleracea, some- 

times severely (Waterhouse 1993b), but too little is known about host specificity or host 

specific strains to evaluate their possible role as classical biological control agents. 

Nevertheless their specificity certainly merits investigation should the need arise. 


There have been no attempts to establish natural enemies as biological control agents for 

P. oleracea. However three insect species have appeared in countries well out of their 

native range, in particular the European weevil Hypurus bertrandi, but also the American 

sawfly Schizocerella pilicornis and the American leaf-mining fly Haplopeodes palliatus. 

These successful, unassisted establishments suggest that there should be little difficulty 

in securing assisted establishments elsewhere. Unfortunately there is no information 

available on what effects, if any, these three insects have had on P. oleracea in their new 

countries, but it is suspected that a group of species may be required to secure substantial 

effects in Australia. 

The sawfly Schizocerella pilicornis appeared in eastern Australia (Queensland and 

New South Wales) in the early 1960s; (Benson 1962, Krombein and Burks 1967) but 

there are no records of it building up in sufficient numbers to cause serious damage. In 

1993 Hypurus bertrandi and Liriomyza caulophaga were bred from P. oleracea leaves in 

Brisbane (R.E. McFadyen pers. comm.), but numbers were too low to cause serious dam- 

age. It is not known whether these species have only become established recently. 

L. caulophaga was previously known only from Beta vulgaris var. cicla (silverbeet) in 

Australia. Larvae tunnel in the soft white spongy tissue between the vascular strands in 

the leaf petioles and midribs and pupate there (Kleinschmidt 1960, 1970, Spencer 1990). 

EGYPT 

Tawfik et al. (1976) recorded Hypurus bertrandi attacking P. oleracea. 

GUAM AND THE NORTHERN MARIANA IS. 

Zimmerman (1957) records Hypurus bertrandi from Guam, Tinian, Saipan and Agrihan, 

some of the specimens from Saipan being taken from the crops of swifts.


Table 4.25.2 Insects restricted to R oleracea or at least to the genus Portulaca (after 

Waterhouse 1993b). 

Species Distribution References 

Coleoptera 

CURCULlONlDAE 

Apion sp. 

Baris arctithorax 

Baris lorata 

Baris portulacae 

Ceutorhynchus oleracae 

Ceutorhynchus portulacae 

Hypurus bertrandi 

Linogeraeus (= Centrinaspis) 

perscitus 

Diptera 

AGROMYZIDAE 

Haplopeodes palliatus 

ANTHOMYIIDAE 

Pegornya dolosa 

CEClDOMYllDAE 

Asphondylia portulacae 

Lasioptera portulacae 

Neolasioptera portulacae 

Lepidoptera 

HELlODlNlDAE 

Heliodines quinqueguttata 

Hymenoptera 

TENTHREDINIDAE 

Schizocerella pilicornis 

Brazil 

Egypt 

Sudan 

India 

Java 

India 

Puerto Rico, 

France, 

Egypt, 

USA, Hawaii, Marianas, 

Australia 

Colombia, Trinidad, USA 

Australia, USA 

Trinidad 

El Salvador, Argentina, 

Colombia, Bolivia, 

Leeward Is, St Kitts 

Nevis, Montserrat, Jamaica 

USA 

Cuba, Florida, St Vincent 

Trinidad, St Kitts Nevis, 

Trinidad,Montserrat 

Puerto Rico 

California, Mexico 

USA, Australia 

Argentina to USA 

d'Araujo e Silva et al. 1968a,b 

Tawfik et al. 1976 

Marshall 19 1 1 

Marshall 19 16 

Marshall 1935 

Marshall 1916 

Wolcott 1948 

Tempsre 1943 

Tawfik et al. 1976 

Clement and Norris 1982 

RE McFadyen pers. comm. 1993, 

Zimmeman 1957 

Bennett and Cruttwell 1972, 

Romm 1937 

R.E. McFadyen pers. comm. 

1993, Romm 1937 

Bennett and Cruttwell 1972 

Cruttwell and Bennett 1972a 

Gagnt 1968, 


Felt 191 1 

GagnC 1968 


Bennett and Cruttwell 1972, 

Cruttwell and Bennett 1972b 

Wolcott 1948 


Krombein and Burks 1967 

Muesebeck et al. 195 1


Table 4.25.3 Natural enemies of Portulaca oleracea: fungi, viruses and nematodes. 


FUNGI 

AIbugo portulacae 

AIbugo portulacearum 

Ascochyta portulacae 

Bipolaris (= Drechslera) 

indica 

Cercospora portulacae 

Cercosporella dominicana 

Dendrographium 

lucknowense 

Dichotomophthora indica 

(= D. lutea) 

Dichotomophthora 

portulacae 

Helminthosporium 

(Bipolaris) portulacae 

Phoma sp. 

Phytophthora palmivora 

Polymyxa betae f. sp. 

portulacae 

VIRUSES 

anemone brown ring 

aster yellows 

beet curly top 

chili vein banding 

clover big vein 



tobacco broad ring spot 

tobacco etch 

tobacco mosaic 

tobacco streak 

NEMATODES 

Criconemella xenoplax 

Ditylenchus dipsaci 

Helicotylenchus indicus 

Helicotylenchus 

multicinctus 

Heterodera glycines 

Europe, Africa, 

Asia, Americas 

Poland 

USSR 

USA 

India 

Dominica 

India 

IMI 1992, Miyanishi & 

Cavers 1980 

IMI 1992 

IMI 1992 

many, including Evans 1987, 

Amaranthus spinosus Kenfield et al. 1989 

IMI 1992 

IMI 1992 

IMI 1992 

USA, West Indies, Helianthus, 

Europe, India Pennisetum 

Europe, Sudan, tarragon, cactus, 

Hawaii, capsicum, 

California, Glycine max 

Jamaica 

Venezuela 

USA Portulaca grandifora 

France tarragon 

Sarawak pepper and several 

weeds 

Bulgaria, Japan sugar beet, wheat, 

many weeds 

Hawaii 

Hawaii 

Hawaii 

Hawaii 

Hawaii 

Bulgaria 

Hungary 

USA 

Malawi 

Hawaii 

Hawaii 

Philippines 

Hawaii 

anemone 

aster 

sugar beet 

chili 

clover 

many economic 

plants 

cucumber, tobacco 

groundnut 

tobacco 

tobacco 

many 

tobacco 

Baudoin 1986, IMI 1992, 

Rao 1966 

Vegh & Le Berre 1984 

Klisiewicz 1985, 

Klisiewicz et al. 1983, 

Mehrlich & Fitzpatrick 1935 

Mitchell 1986 

IMI 1992, Rader 1948, 

Strider & Chi 1984 

Vegh & Le Berre 1984 

Anon 1979 

Abe & Ui 1986, 

Vrbanov & Krumov 1989 






Dikova 1989, 

Nasser & Basky 1988, 

Dodds & Taylor 1980 

Adams 1967 



Eugenio & del Rosario 1962 


USA many legumes and Zehr et al. 1990 

other plants 

USSR polyphagous Kholod 1983 

India polyphagous Rahman & Khan 1986 

Brazil, Ivory banana Luc et al. 1990, 

Coast Zem & Lordello 1983 

Colombia soybean Quintero et al. 1988 

I (continued on next page)



Heterodera rnarioni 

Hoplolairnus indicus 



Meloidogyne hapla 



Pratylenchus rninutus 

Pratylenchus sp. 

Radopholus sirnilis 

Rotylenchulus reniforrnis 

Tylenchorhynchus 

brassicae 

Hawaii 

USA 

Cuba 

USA 

Hungary 

India, USA, 

Philippines 

India 

Hawaii 

Ivory Coast 

Ivory Coast 

India, USA, 

Hawaii 

India 

eggplant, tomato 

coffee 

tobacco 

several weeds 

polyphagous 

polyphagous 

Musa sp., 

several weeds 

ornamentals and 

many weeds 

polyphagous 

Linford & Yap 1940 

Rahman & Khan 1986 

Izquierdo et al. 1987 

Tedford & Fortnum 1988 

Dabaj & Jenser 1990 

Maqbool et al. 1986, 

Tedford & Fortnum 1988, 

Valdez 1968 

Maqbool et al. 1986 




Heald et al. 1974, 

Inserra et al. 1989 

Khan & Khan 1985 

Linford & Yap 1940 

Rahman & Khan 1986 

HAWAII 

Pigweed was established in Hawaii prior to 187 1 (Hillebrand 1888). Hypurus bertrandi, 

originally misidentified as Ceutorhynchus sp., was reported in 1950 to be numerous 

enough in many cases to defoliate P. oleracea and to cause it to collapse as if sprayed 

with a herbicide (Bianchi 1955). Nevertheless, in 1992, Hawaiian weed scientists considered 

it as one of their worst weeds (W.C. Mitchell pers. comm. 1992), so the control 

exerted by H. bertrandi and v'arious non-specific insects is clearly insufficient. 

Zimmerman (1957) postulates that H. bertrandi was introduced from the Marianas to 

Hawaii, possibly with war material being returned from the battlefields. However it is 

more likely to have moved in the reverse direction. 

INDIA 

Zaka-ur-rab (1991) records Hypurus bertrandi as one of the leaf-mining weevils of the 

Indian subcontinent. 

PUERTO RlCO 

Wolcott (1948) recorded H. bertrandi from Puerto Rico, but no other information is 

available. 

Major Natural Enemies 

Summarised below is what is known of the biology of nine of the natural enemies listed 

in table 4.25.2. 

Apion sp. Coleoptera: Curculionidae 

Apion sp. produces galls in the flower buds of P. oleracea in Brazil (d'Araujo e Silva et 

al. 1968a,b) and Apion larvae also cause significant damage by gall formation in flowers 

in northern Argentina (Bennett and Cruttwell 1972, Bennett pers. comm. 1992).


Asphondylia portulacae Diptera: Cecidomyiidae 

Oviposition by this flower gall midge into the very small pigweed buds causes them to 

develop abnormally. Usually only one larva develops per bud and occupies a chamber in 

the swollen receptacle. Galled flowers do not produce seed. A. portulacae is heavily 

attacked by parasitoids (Bennett and Cruttwell 1972) and might be an important natural 

enemy if freed from them. The genus Asphondylia is considered to be highly host specif- 

ic and 52 of its 54 species are known from only a single host. Each of the two exceptions 

only attacks two plants of the same genus and Bennett and Cruttwell (1972) suggested, 

on the basis of this information, that host specificity testing was unnecessary. 

Baris arctithorax Coleoptera: Curculionidae 

In Egypt this weevil forms stem galls on pigweed, but is not known from any economic 

plant. Eggs are laid singly in cavities gnawed in the stem by the female, leading to the 

production of single closed galls in which the larvae feed. Pupation occurs in the soil. 

Young infested plants produce weak vegetative growth, few seeds and may even be 

killed. Adult weevils feed on the surface of leaves causing white blotches or holes. At 

temperatures from 25 to 30°C the development time from egg to adult is about 40 days. 

Infestations of up to 74% of plants are recorded in summer and a peak of 95% in autumn 

(Awadallah et al. 1976, Tawfik et al. 1976). 

Haplopeodes palliatus Diptera: Agromyzidae 

The genus Haplopeodes contains 13 species, all from the Americas and known on only 

four plant families-Portulacaceae (1 species) Amaranthaceae (3 species), 

Chenopodiaceae (2 species) and Solanaceae (8 species) (Spencer 1990). Each appears to 

be specific to a single genus and H. palliatus is known only from l? oleracea. It is a typi- 

cal leaf miner. 

Heliodines quinqueguttata Lepidoptera: Heliodinidae 

Eggs, which are laid singly or in groups of up to 6, hatch in 5 to 6 days and larvae wan- 

der some distance over the plant before mining into a leaf, stem or seed capsule. After 7 

to 8 days the fifth instar larva leaves the mine and pupates within a flimsy silk cocoon 

attached to the stems or leaves of the plant. Larvae are attacked by a braconid endopara- 

sitoid, Pholetesor (= Apanteles) sp. (circumscriptus group). 

Host specificity tests were carried out on a wide variety of economic and non- 

economic plants, but development was completed only on Portulaca oleracea, P. pilosa 

(also weedy) and the ornamental P. grandiflora. However, in the field in Trinidad, neither 

P. pilosa nor P. quadriJida (also weedy) were attacked and l? grandiflora was not grown. 

There appear to be no records of Heliodines species attacking crops and each species 

appears to be restricted to a single plant family. Cruttwell and Bennett (1972b) suggested, 

therefore, that it should be considered as a biological control agent. 

Hypurus bertrandi Coleoptera: Curculionidae 

This tiny (2mm long) weevil has spread unaided from its native France to Egypt (prior to 

1926) (Hoffman and Tempbre 1944, Tawfik et al. 1976), Puerto Rico (Wolcott 1948), 

Hawaii (1950) (Davis 1955, Maehler 1954), Guam and the Northern Marianas


(Zimmerman 1957), California (1980) and Queensland (1993) (R.E. McFadyen pers. 

comm.). 

Eggs are deposited singly and larvae mine the leaves. Infested leaves wilt and fall; 

and the larvae then migrate to fresh leaves, each destroying four or five in its lifetime. If 

no undamaged leaves are available the outer tissues of stems are attacked. Pupation 

occurs in a cell formed by soil particles cemented by fecal secretion and, in France, 

adults overwinter under the bark of trees. Adults feed on leaf margins, stems and devel- 

oping seed capsules. Development is rapid, from egg to adult in 10 days at 32.2OC and 

under 16 hours light. Z? oleracea is its only reported host plant. In France it is parasitised 

by a number of wasps (Tawfik et al. 1976, Clement and Norris 1982, Hoffmann and 

Tempbre 1944, Norris 1 985, Tempbre 1943, 1 944, 1950). 

Neolasioptera portulacae Diptera: Cecidomyiidae 

Oviposition in the stem by this midge leads to globular galls up to 1.5 cm in diameter, 

each containing up to 10 larvae. Galls retard or prevent growth and also flower and seed 

production. In open, infertile sites every pigweed stem may be infested but, in vigorous 

growth or in shaded sites, the level of attack is usually very low. The larvae are heavily 

attacked by parasitoids. 

All except one of the 51 species of Neolasioptera are restricted to one plant genus 

and the remaining species only attacks two plant genera. This was taken by Bennett and 

Cruttwell (1972) to indicate that N. portulacae is sufficiently host specific to be 

employed for biological control. 

Pegomya dolosa Diptera: Anthomyiidae 

Eggs are laid singly on the underside of the pigweed leaf and hatch after about 3 days. 

The larvae are leaf miners, devouring the contents of the leaf and then leaving to enter 

another. Two or more leaves are commonly destroyed by each larva. After feeding for 

about 7 days larvae leave to pupate in the soil, later emerging as 3 to 4 mm long adults. 

Eggs are parasitised and larvae are attacked by a pteromalid wasp. 

Schizocerella pilicornis Hymenoptera: Tenthredinidae 

This leaf-mining sawfly occurs naturally over a very wide range from Argentina and 

Brazil to USA (Muesebeck et al. 195 1). It appeared unaided in eastern Australia (Benson 

1962, Krombein and Burks 1967). There are two biotypes. The larvae of one which is 

widespread in USA mines the leaves, whereas those of the other (from Mississippi north- 

wards) feeds externally on the leaves. 

Females mate soon after emergence and lay up to 40 eggs singly in the edges of the 

leaves. The larvae mine the leaves, damaging each to the point of collapse before moving 

to another. At least two leaves are destroyed by each larva. The mature larvae enter the 

soil and spin cocoons. The life cycle can be completed in 13 days and there are a number 

of generations each year (Clement and Norris 1982, Gorske et al. 1976). In California 

prepupae in diapause overwinter in the soil. Adults live for a day and do not feed. In 

California up to 84% of Z? oleracea leaves were severely damaged, leading to defoliation 

and sometimes death of the plant. When Z? oleracea was protected by insecticide from 

both S. pilicornis and Hypurus bertrandi it produced about four times as much seed as

218 


unprotected plants, although the latter still produced enough (4000 to 5000/m2/day) to 

maintain a high seed bank in the soil (Force 1965, Garlick 1922, Gomes de Lima 1968, 

Gorske et al. 1977, Norris 1985, Webster and Mally 1900). 

S. pilicornis has not been recorded from any economic plant and, in starvation tests, 

was only able to feed on P. oleracea and the related Montia perfoliata (Gorske et al. 

1976). A microsporidan, Nosema pilicornis causes high mortality in infected S. pilicornis 

larvae in USA and should be eliminated during any transfer of the sawfly to new areas 

(Gorske and Maddox 1978). 

Comment 

It is generally believed that, through coevolution, there are likely to be a number of spe- 

cific (or nearly specific) natural enemies of an organism in its area of origin. 

Furthermore, that not all such organisms will have accompanied their host when it has 

moved outside its area of origin. If this generalisation is applied to P. oleracea it can be 

seen that 8 of its 14 specific (or nearly specific) natural enemies are of American origin 

but no more than 2 from any other region of the world (Table 4.25.2). The inference from 

this is that P. oleracea is probably of American origin, an inference supported by the 

finding of seeds in Louisiana, Illinois and Kentucky dating between lOOOBC and 750AD 

and pollen and seeds in Ontario sediments dating back to 1350AD (Miyanishi and Cavers 

1980). This suggests that, if it is desired to evaluate insects additional to those listed in 

table 4.25.2, they should first be sought from the Americas and possibly from amongst 

those listed by Waterhouse (1993b). 

The family Portulacaceae is relatively small with 20 genera and about 250 species 

worldwide, of which the genus Portulaca contains about half (West 1990). Very few 

Portulacaceae are cultivated: Portulaca grandiflora as a brightly flowering garden plant, 

Talinum paniculatum and T. triangulare as pot herbs (but they are sometimes regarded as 

weeds), Montia fontana for salads, Lewisia spp. as rockery plants and Anacampseros as a 

succulent; but these are not considered to be of great economic importance (Cruttwell 

and Bennett 1972a). 

From what is known about the biology of the insects in table 4.25.2 it appears that, 

if a group of them is established in a new country without their own natural enemies, 

they should cause serious damage to l? oleracea and significantly reduce its competitive- 

ness and seed production.



(after Holm eta/. 1977)

Map 4.26 Rottboellia cochinchinensis 

4.26 Rottboellia cochinchinensis 221 


Rottboellia cochinchinensis is probably of Southeast Asian or Indian origin, although one 

biotype may have evolved in East Africa. Very little is known of its insect enemies in 

these regions and they certainly merit investigation. Current research on fungal 

pathogens indicates that Sporisorium ophiuri is a potential agent for the Americas and 

other regions where it does not yet occur. 

If preliminary surveys for specific insect enemies in Asia and Africa support a 

Southeast Asian origin it is unlikely to be an attractive early target for classical biological 

control in this region.


4.26 Rottboellia cochinchinensis (Lour.) W.D. Clayton 

(= Rottboellia exaltata) 

Poaceae 

itch grass; myet ya, myet ya nge (Myanmar), branjangan (Indonesia), aguitigay 

(Philippines), yaa prong khaai (Thailand) 

Rating 

+++ Indo, Phil 

12 ++ Thai 

+ Myan, Laos, Viet, Msia 

Origin 

Uncertain but probably India to Southeast Asia; one biotype in East Africa. 

Distribution 

Worldwide, and weedy between 23' north and south latitudes (Holm et al. 1977). 

Formerly known as Rottboellia exaltata. Two biotypes have been recognised in USA and 

at least five in the Philippines (Fisher et al. 1987). The origin of R. cochinchinensis has 

not been completely resolved. It is said to be native to India (Holm et al. 1977), but in his 

survey of fungal pathogens and their specificity (or lack of it) Evans (pers. comm. 1992) 

was led to the conclusion that it may be Southeast Asia; and furthermore, that there is a 

distinct East African biotype, with possibly specific natural enemies of its own. It is 

interesting that, of the 14 Kenyan tribes visited by Ellison and Evans (1990), 13 had a 

separate and specific vernacular name for itch grass. Although it was a common weed 

and dense stands occurred at the edge of fields and along roadsides, control was achieved 

by early season hoeing and none of the farmers suggested it was a major problem. This 

evidence was taken as reinforcing the theory that East Africa is the centre of origin at 

least of the local biotype of the weed. 


R. cochinchinensis is a tall, erect, strongly tufted, annual grass growing to 3 m. It has stilt 

roots. Its leaves and stems have long, sharp, fragile, siliceous, irritating hairs that break 

off in the flesh on contact. The inflorescence is a single cylindrical spike. 

Importance 

It is an aggressive C4, annual grass of 18 tropical and subtropical crops in 28 countries, 

including maize, rice, sorghum, soybeans and sugarcane. The heaviest infestations occur 

in the Caribbean, Central America, and parts of South America, to which it is a relatively 

recent introduction; also a widespread weed in southern Africa. It is an important weed in 

sugarcane, maize and upland rice in the Philippines. It is often a primary coloniser of dis- 

turbed land. It flowers all year round. Reproduction is by seed and up to 8000 may be


produced per plant. The seed is about the size of a rice grain and is not easily separated 

from intermingled rice grains. Some seeds germinate immediately, whereas others lie 

dormant for varying periods, sometimes years. Buried seeds may germinate and emer- 

gence take place from a depth of 15 cm. It is common in open, well-drained places, but 

also grows in wet places and even in shallow water. It commonly occurs on contour 

banks and roadsides. It prefers sunny or no more than moderately shaded situations, but 

can grow in deep shade. Its many needle-like hairs deter hand removal of older plants, 

since the hairs penetrate hands and clothing and result in painful infections. 

R. cochinchinensis is sometimes grazed and used for green fodder, although avoided 

at times by some animals because of its sharp hairs. 


Fungi are the only natural enemies (Tables 4.26.1 to 4.26.3) for which there is any rea- 

sonably comprehensive information. Fungi have been surveyed and their specificity is 

being examined in a joint International Institute of Biological Control and Long Ashton 

Research Station project covering East Africa, South America, India, Nepal, Sri Lanka 

and Thailand (Ellison 1992, Ellison and Evans 1990, 1993, Evans 1991, Natural 

Resources Institute 1992). 

As indicated earlier, R. cochinchinensis shows high biotype variation between coun- 

tries and this is correlated with varying levels of susceptibility to different fungal isolates. 

Also, a clear positive correlation was found between high pathogen virulence and inade- 

quate specificity to itch grass (Ellison 1992). Maize (Zea mays) proved to be the crop 

species most at risk from itch grass pathogens, which is not surprising in view of the 

close evolutionary relationship of the two genera involved. This suggests that the use of 

fungi for classical biological control of itch grass may not show great promise, but that 

their use as mycoherbicides might prove effective. All except one of the fungi from tropi- 

cal America that have been tested are non-specific to itch grass and most are local 

pathogens that have transferred from local grasses (Evans 1987). In Kenya, in addition to 

at least 10 non-specific fungi attacking itch grass, a head smut, Sporisorium 

(= Sphacelotheca) ophiuri was found, which appears to be restricted to Rottboellia and 

the closely related genus Chasmopodium (Ellison and Evans 1990, Zundel 1953). 

S. ophiuri is recorded as occurring in East Africa, Sri Lanka, Philippines and 

Thailand, but apparently not in the Americas. It is often locally damaging, significantly 

reducing vigour and virtually eliminating seeding. Its host specificity is under detailed 

investigation (Ellison and Evans 1993, Evans 1991) as a potential candidate for classical 

biological control for areas where it does not already occur. In an annual weed where 

seeds are the only means of propagation, a destructive seed head disease, such as S. ophi- 

uri, is a highly promising biological control agent (Evans 1991). 

A Curvularia isolate from Trinidad proved highly damaging to itch grass, while not 

damaging rice, maize, sugarcane or pearl millet (Evans 1991). A Curvularia from 

Somalia was able to kill R. cochinchinensis in a few days, but was also able to infest 

maize. However the crop readily recovered (Ellison 1992). If further tests confirm its 

specificity, it may be a potential biological control agent. The same applies to Puccinia


rottboelliae about which less is recorded (Evans 1987). Special attention is now being 

paid to the possibility of developing preparations of one or more of these fungi as a 

mycoherbicide. An isolate from Thailand of Colletotrichium sp. which appears to be spe- 

cific to itch grass has been selected from 900 fungal samples and field trials have already 

demonstrated that an appropriate formulation has potential against this weed, particularly 

when combined with low doses of herbicide (Ellison 1992, Ellison and Evans 1993, 

Natural Resources Institute 1992). 

Surprisingly few insects (Table 4.26.1) have been recorded attacking itch grass and 

only one unidentified gall midge recorded in India from Rottboellia compressa (Barnes 

1946) might, if it attacks R. cochinchinensis also, be specific enough to be a candidate 

agent. It is regrettable that parallel observations were not made on insects during the 

extensive fungal surveys. In East Africa a stem borer, a lepidopteran leaf feeder and a fly 

larva all proved to be non-specific graminaceous feeders (H.C. Evans pers. comm. 1992, 

1993). 

R. cochinchinensis is an alternative host for a number of viruses, almost all of them 

serious diseases of maize (Table 4.26.3). It is surprising that the only record encountered 

dealing with nematodes related to a study of 16 plant parasitic species attacking sugar- 

cane in the Philippines. This found that itch grass was not infected by Meloidogyne sp. 

(Reyes and Beguico 1978). 

Comment 

Rottboellia belongs to the same grass tribe (Andropogoneae), but not to the same sub- 

tribes, as Saccharum, Sorghum and Zea (Table 4.26.4). This suggests that candidates for 

classical biological control of this weed will have to pass extensive host specificity test- 

ing against all of the crop and pasture grasses belonging to these and related genera 

before being considered for release. 

Table 4.26.1 Natural enemies of Rottboelliu cochinchinensis: insects. 


Orthoptera 

GRYLLIDAE 

Euscyrtus concinnus Philippines Eleusine indica, Barrion & Litsinger 1980 

Dactylocteniurn 

aegyptiurn, Cyperus 

rotundus, Digitaria 

sanguinalis 

Pteronernobius sp. Sierra Leone Alghali & Domingo 1982 

Hemiptera 

APHlDlDAE 

Myzus persicae Peru a very wide range Ortiz 198 1 

Uroleucon (= Dactynotus) 

. ambrosiae USA sorghum, maize Koike 1977 

I 




CICADELLIDAE 

Nephotettix nigrornaculatus Sierra Leone 

DELPHACIDAE 

Peregrinus rnaidis Venezuela 

Coleoptera 

COCCINELLIDAE 

Chnootriba (= Epilachna) 

similis Sierra Leone 

Diptera 

AGROMYZIDAE 

Pseudonapornyza 

philippinensis Philippines 

CEClDOMYllDAE 

a gall midge India 

MUSCIDAE 

Atherigona soccata Kenya 

Lepidoptera 

LYMANTRIIDAE 

Psalis pennatula Kenya 

NOCTUIDAE 

Sesarnia sp. Ghana 

Spodoptera frugiperda USA 

PYRALIDAE 

Chilo sp. 

sp.? 

Ghana 

Kenya 

rice, Ischaernurn Alghali & Domingo 1982 

rugosurn, Paspalurn 

vaginaturn 

a virus transmitter Ferreira et al. 1989, Migliori 

on many hosts & Lastra 1980, 198 1, 

Trujillo et al. 1974 

Alghali & Domingo 1982 

known only from Spencer 1961 

R. cochinchinensis 

recorded only from Barnes 1946 

Rottboellia cornpressa 

rice Ogwaro 1978 

generalist Poaceae H.C. Evans pers. comm. 1993 

leaf eater 

Sampson & Kumar 1986 

many graminaceous Rajapakse et al. 1988 

crops 

Sampson & Kumar 1986 

generalist Poaceae H.C. Evans pers. comm. 1993 

leaf eater 

Table 4.26.2 Natural enemies of Rottboellia cochinchinensis: fungi. 

Fungi Location Other hosts References 

Ascochyta sp. 

Bipolai-is perotidis 

Cercospora spp. 

Kenya no tests on other Ellison & Evans 1990, 1993 

plants 

Australia many other grasses QDPI, unpublished 

Kenya, Ethiopia, maize mildly Ellison & Evans 1990, 1993 

Zanzibar, 

Madagascar, 

Americas, SE Asia 




Fungi 

Cercospora fusimaculans 

Cercospora rottboelliae 

Cochliobolus 

(Helminthosporiurn) bicolor 

Cochliobolus (Curvularia) 

cymbopogonis 

Cochliobolus heterostrophus 

(Drechslera maydis) 

Colletotrichium sp. 

Coniothyrium sp. 

Curvularia spp. (many) 

Diaporthe (Phomopsis) sp. 

Diplodia sp. 

Fusarium moniliforme 

(Gibberella fujikuroi) 

Glornerella (Colletotrichium) 

grarninicola 

Leptosphaeria sp. 

Magnaporthe (Pyricularia) 

grisea 

Phaeoseptoria sp. 

Phyllachora sacchari 

Puccinia rottboelliae 

Location Other hosts References 

Sudan, Zambia, many grasses 

Ghana, Guinea, 

Togo,.Uganda, 

Jamaica 

Guinea 

Zimbabwe, sugarcane, maize, 

Somalia pearl millet 

Bolivia, Kenya, sugarcane, maize, 

Trinidad, pearl millet, 

Zanzibar, USA, sorghum 

SE Asia 

Kenya, Papua sugarcane, maize 

New Guinea, 

SE Asia 

Thailand (this species is close 

to C. graminicola) 

Africa, SE Asia 

Trinidad, Papua most (not all) attack 

New Guinea, economic crops 

Madagascar, 

Somalia, Zanzibar 

Kenya, SE Asia maize mildly 

Kenya no tests on other 

plants 

Guatemala the particular strain 

tested had limited 

host range with no 

symptoms in maize, 

sorghum or sugar 

Jimenez et al. 1990 

cane, but other strains 

attack these and rice 

India, Nepal, No Colletotrichiurn Evans 1987, 1991 

Sri Lanka, infection observed 

Thailand in East Africa 

Kenya, SE Asia no tests on other 

plants 

Ellison & Evans 1990, 1993 

Kenya, maize mildly, also 

Zimbabwe Eleusine spp. 

Kenya, SE Asia no tests on other 

plants 

Asia, Nigeria, 

Sicily, Argentina 

Kenya, Mada- limited host range 

gascar, Ghana, 

Uganda, Zambia, 

Ethiopia, Guinea, 

Nigeria, Sierra 

Leone, Sudan, 

Zimbabwe, India 

Evans 1987 

Evans 1987 

Ellison & Evans 1990. 1993 

Ellison & Evans 1990, 1993, 

Evans 1987, Walker & 

White 1979 


Evans 1987 

Ellison 1992, Ellison & Evans 

1993 

Ellison & Evans 1993 

Ellison 1992, Evans 1991 


Ellison & Evans 1990. 1993 


Evans 1987 


Anahosur and Sivanesan 1978 


Evans 1987 



Fungi Location Other hosts References 

Pyrenochaeta sp. SE Asia 

Sphacelotheca rottboelliae Malawi, India 

Sporisorium (= Sphacelotheca) Kenya, Somalia, 

ophiuri Sudan, Uganda, 

Zimbabwe, 

Sierra Leone, 

Sri Lanka, 

Philippines 

Ustilago scitaminea Philippines 

Ellison & Evans 1993 

also on Saccharum Evans 1987 

spontaneum 

limited host range, Ellison & Evans 1990, 1993, 

and not present Evans 1987, 1991 

in the Americas 

Latiza 1980 

Table 4.26.3 Natural enemies of Rottboellia cochinchinensis: viruses. 

Virus Location Other hosts References 

corn leaf gall virus Philippines Agati & Calica 1950 

maize stripe tenuivirus USA sorghum Bradfute & Tsai 1990 

maize stripe virus USA sorghum Gingery et al. 1981 

maize hoja blanca Venezuela sorghum Ferreira et al. 1989 

maize white leaf Venezuela Trujillo et al. 1974 

maize rayado fine Texas several grasses Nault et al. 1980 

maize mosaic Guadeloupe, 

French Guinea sorghum Migliori & Lastra 198 1 

maize dwarf mosaic USA sorghum, sugarcane Gillespie & Koike I973 

maize yellow mottle Nigeria Thottapilly et al. 1992 

virus like disease of maize Guadaloupe sorghum Migliori & Lastra 1980 

rice leaf gall Philippines Agati & Calica 1950 

Table 4.26.4 Taxonomic position of the major exotic grass weeds (bold text) in rela- 

tion to the major genera of crops in the family Poaceae (= Gramineae). 

Subfamily Tribe Sub-tribe Genera 

Bambusoideae Bambuseae 

Oryzeae 

Pooideae Triticeae 

Aveneae 

Chloridoideae Eragrostideae 

Cynodonteae 

Panicoideae Paniceae 

Andropogoneae 

Maydeae 

Bambusa 

Oryza 

Hordeum, Secale, 

Triticum 

Avena 

Eleusine, Eragrostis 

Cynodon 

Setariinae Echinochloa, Panicum, 

Paspalum, Setaria 

Digitariinae Digitaria 

Cenchrinae Pennisetum 

Saccharinae Saccharum 

Sorghinae Sorghum 

Rottboelliinae Rottboellia 

Tripsacinae Zea




Map 4.27 Sphenoclea zeylanica 

4.27 Sphenoclea zeylanica 229 


Goose weed, Sphenoclea zeyhnica, is native to Tropical Africa where it is not regarded 

as a weed, although there are no natural enemies recorded from it there. In Southeast 

Asia, where it is an important weed of rice, and in India, it is sometimes severely affected 

by a fungus which may have some promise as a bioherbicide. 

A survey for natural enemies in its area of origin would be required to evaluate 

the prospects for its biological control.


4.27 Sphenoclea zeylanica Gaertn. 

Sphenocleaceae 

gooseweed; goenda (Indonesia), silisilihan (Philippines), pakpawd, phak pot 

(Thailand) xa b6ng (Vietnam) 

Rating 

+++ Msia 

14 ++ Thai, Camb, Phil 

+ Myan, Laos, Viet, Brun, Indo 

Origin 

Tropical Africa 

Distribution 

In tropical and subtropical regions across the world. From Iran extending eastwards to 

Indonesia and the Philippines, also China, Japan, USA, the ~aribbean, Guyana, Surinam 

and Madagascar. Not reported from Papua New Guinea, Australia or the oceanic Pacific. 


S. zeylanica is an erect, fleshy, herbaceous annual, often with much branched, hollow 

stems, growing to 1.5 m; leaves alternate, oblong to lanceolate, tapering to both ends; 

flowers sessile in dense spikes, terminal, whitish; seed yellowish brown, 0.5 mm long; 

roots cord-like. 

Importance 

S. zeylanica is unusual in that it is not reported as a weed in any crop except rice (Holm 

et al. 1977). It thrives in damp ground at altitudes up to 350 m. In Africa it grows in the 

mud of tidal creeks, but does not have this habit in Malaysia. It occurs on the sides of 

ponds and along ditches and rivers, on dry river beds and in seasonal swamps. It prefers 

stagnant water sites. It reproduces continuously by seed in the Philippines. In spite of its 

competition with the rice plant, S. zeylanica can give valuable practical control (up to 

99%) of populations of rice nematodes (Hirschmaniella spp.), with the additional benefit 

of increased soil nitrogen. It acts through the production of toxic plant exudates 

(Mohandes et al. 1981). 

In Java, young plants and tips of older plants are steamed and eaten with rice. 


The only natural enemy encountered in the literature search is a fungus (Table 4.27.1). 

This was a severe infestation of the fungus Cercosporidiurn helleri on the lower surface 

of S. zeylanica leaves in India (Ponnappa 1967). The affected leaves became deformed 

and fell off. Similar fungi capable of causing death of the weed were observed at Los

4.27 Sphenoclea zeylanica 231 

Banos, Philippines and at Prey Phadu, Cambodia (Moody et al. 1987). If this fungus 

proves to be adequately specific it may have some value as a bioherbicide. 

Table 4.27.1 Natural enemies of Sphenoclea zeylanica 


NEMATODE 

Meloidogyne graminicola Luc et al. 1990 

FUNGUS 

Cercosporidium helleri Cambodia, India, Moody et al. 1987, 

Philippines Ponnappa 1967

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6 Index of scientific names of insects 

abdominalis, 


Acanthoscelides pigricola 

Kingsolver Col.: 

Bruchidae 152 

Acanthoscelides puniceus 

Johnson Col.: Bruchidae 

150,152,153 


quadridentatus (Schaeffer) 

Col.: Bruchidae 150, 152, 

153 

Acanthoscelides zebratus 

Kingsolver Col.: 

Bruchidae 152 

Achaea janata (Linnaeus) 

Lep.: Noctuidae 105 

Acigona infusella, see 

Haimbachia infusella 

Acrida turrita Linnaeus Ort.: 

Acrididae 133 

Acrosternum herbidum (Stiil) 

Hem.: Pentatomidae 144 

Actinote anteas (Doubleday 

and Hewitson) Lep.: 

Nymphalidae 39,44,47, 

48 

aculeatum, Coelocephalapion 

acuminata, Paulinia 

acuta, Leptocorisa 

Adaina bipuncta Moschler 

Lep.: Pterophoridae 129 

adipalis, Ategumia 

aeneicollis, Horismenus 

aerea, Colaspis 

aerea, Maecolaspis 

Aerenica hirticornis (Klug) 

Col.: Cerambycidae 40 

afinis, Neohydronomus 

agamemnon, Chi10 

Agathis Hyrn.: Braconidae 

206 

Agraulis vanillae (Linnaeus) 

Lep.: Nymphalidae 187, 

188 

Agrmyza parvicornis Loew 

Dip.: Agromyzidae 64 

Agromyza proxima Spencer 


Agrotera basinotata 

Harnpson Lep.: 

Pyralidae 122 

Agrotis ipsilon (Hufnagel) 

Lep.: Noctuidae 21 2 

albicinctus, Orosius 

albiguttalis, Sameodes 

Alcidodes Col.: Curculionidae 

1 22 

aliphera, Eueides 

allecta, Calycomyza 

Altica cyanea (Weber) Col.: 

Chrysomelidae 121, 1 22 

Amata huebneri Boisduval 

Lep.: Amatidae 134 

Amauromyza Dip.: 

Agromyzidae 32,57 

Amauromyza maculosa 

(Malloch) Dip.: 


amazonus, Longitarsus 

arnbrosiae, Dactynotus 

ambrosiae, Uroleucon 

Ammalo arravica, see 

Pareuchaetes 


Ammalo insulata, see 


Amsacta gangis, see 

Creatonotos gangis 

Amsacta lactinea (Cramer) 

Lep.: Arctiidae 45 

anaphalidis, Masonaphis 

anaphalidis, Neomasonaphis 

andropogonis, Dyodiplosis 

andropogonis, Orseolia 

anisodactylus, Sphenarches 

Anoecia querci , see 

Stegophylla querci 

antem, Actinote 

antica, Pochazia 

antonii, Helopeltis 

Apanteles Hyrn.: Braconidae 

206 

Apanteles cordoi De Santis 

Hyrn.: Braconidae 80 

Apanteles creatonoti Viereck 

Hyrn.: Braconidae 43, 

Apantelesfluitantis De Santis 

Hyrn.: Braconidae 80 

Aphis Hem.: Aphididae 121 

Aphis citricola, see Aphis 

spiraecola 

Aphis coreopsidis (Thomas) 

Hem.: Aphididae 30 

Aphis craccivora Koch Hem.: 

Aphididae 13,45, 105 

Aphis fabae Scopoli Hem.: 

Aphididae 41,64, 188 

Aphis gossypii Glover Hem.: 

Aphididae 13,45, 105, 133 

188 

Aphis illinoisensis Shimer 


Aphis nigricauda, see Aphis 

spiraecola 

Aphis spiraecola Patch Hem.: 

Aphididae 1 3,41,44,45, 

133,188 

aphodoides, Temnodachrys 

Apion Col.: Apionidae 130, 

213,215 

Apion brunneonigrum 

BCguin-Billecocq Col.: 

Apionidae 38,39,42-44, 

46-48 

Apion luteirostre Gerstaecker 

Col.: Apionidae 30, 127, 

129,130 

Apsilops sp. Hyrn.: 

Ichneumonidae 206 

archboldi, Liriomyza 

Archips Lep.: Tortricidae 

23 

Archips micaceana (Walker) 

Meyrick Lep.: Tortricidae 

123 

arctithorax, Baris 

argentiniensis, Disonycha 

Argentinorhynchus bennetti 

O'Brien and Wibmer Col.: 

Curculionidae 200

278 Biological Control ( ~f Weeds: Southeast Asian Prc 

Argentinorhynchus breyeri 

Br2thes Col.: 

Curculionidae 200 

Argentinorhynchus bruchi 

(Hustache) Col.: 

Curculionidae 200,204 

Argentinorhynchus minimus 



Argentinorhynchus 

squamosus (Hustache) 

Col.: Curculionidae 200 

argentinus, Horciacinus 

Argyractis subornata 

(Hampson) Lep.: Pyralidae 

72,201 

Argyresthia leuculias 

Meyrick Lep.: 

Argyresthiidae 122 

armatus, Chariesterus 

armigera, Dicladispa 

Aroga Lep.: Gelechiidae 15 1 

arravaca, Ammalo 

arravaca, Pareuchaetes 

artemisia, Dynamine 

Arthrocnodax meridionalis 

Felt Dip.: Cecidomyiidae 

48 

Ascia buniae phaloe, see 

Perrhybris phaloe 

aspersa, Sibinia 

asperulus, Ceutorhynchus 

Asphondylia Dip.: 

Cecidomyiidae 2 1 6 

Asphondylia bidens 

Johannsen Dip.: 

Cecidomyiidae 3 1 

Asphondylia corbulae Mohn 

Dip.: Cecidomyiidae 40 

Asphondylia portulacae 

Mohn Dip.: Cecidomyiidae 

213,216 

Astylus lineatus (Fabricius) 

Col.: Melyridae 22 

Asynonychus godmani Crotch 


Ategumia adipalis (Zeller) 

Lep.: Pyralidae 120, 122, 

123 

Ategumia fatualis (Lederer) 


123 

Atherigona soccata Rondani 

Dip.: Muscidae 225 

atilus, Sphaeniscus 

atricornis, Phytomyza 

atrovenosa, Nisia 

Atractomorpha crenulata 

(Fabricius) Ort.: Acrididae 

78 

Aulacorthum solani 

(Kaltenbach) Hem. 

Aphididae 13 

Aulacochlamys Col. 

Chrysomelidae 39,49 

aurata aurantior, 

Pareuchaetes 

aurata aurata, Pareuchaetes 

aurata, Pareuchaetes 

Austelasmus Hym.: 

Elasmidae 5 1 

Autoba versicolor (Walker) 


avenae, Sitobion 

Azarma densa, see Bellura 

densa 

azia, Thecla 

azia, Tmolus 

Bactrocera dorsalis (Hendel) 

Dip.: Tephritidae 120, 122 

Bactrocera pedestris, see 

Bactrocera dorsalis 

bada, Parnara 

Baliothrips biformis, see 

Stenchaetothrips biformis 

baridiiformis, 

Psuedoderelomus 

Baris Col.: Curculionidae 29, 

30,40 

Baris arctithorax (Pic) Col.: 

Curculionidae 2 12,213, 

216 

Baris lorata Marshall Col.: 

Curculionidae 2 1 3 

Baris portulacae Marshall 

Col.: Curculionidae 2 13 

basalis, Xyonysius 

basinotata, Agrotera 

basui, Tetraneura 

Bellura densa (Walker) Lep.: 

Noctuidae 73,79,80 

Bemisia costa-limai, see 


Bemisia tabaci (Gennadius) 

Hem.: Aleyrodidae 14, 16, 

99,100,105 

bennetri, Argentinorhynchus 

bertrandi, Hypurus 

bessus, Nisoniades 

bibiana, Syphrea 

biblis, Didonis 

bicincta fraterna, 

Monecphora 

bicincta fraterna, Prosapia 

bicolor, Stirellus 

bidens, Asphondylia 

bidentis, Melanogromyza 

bifidalis, Loxostege 

biformis, Baliothrips 

biformis, Stenchaetothrips 

bigaria, Desmogramma 

binaria, Micrutalis 

bipartista, Dactylispa 

bipuncta, Adaina 

bissellata, Coelophora 

Blissus leucopterus (Say) 

Hem.: Lygaeidae 90 

botanephaga, Sesamia 

Bothrogonia ferrugenea 

(Fabricius) Hem.: 

Cicadellidae 133 

Brachycaudus helichrysi 

(Kaltenbach) Hem.: 

Aphididae 13 

Brachycyttarus griseus De 

Joannis Lep.: Psychidae 

181,182 

Brachymeria ovata (Say) 

Hym.: Chalcididae 187 

Brachymeria russelli Burks 

Hym.: Chalcididae 130 

breyeri, Argentinorhynchus 

brickelliae, Neolasioptera 

bruchi, Argentinorhynchus 

bruchi, Neochetina 

bruchi, Ochetina 

brunella, Selca 

brunnea, Myrmicaria 

brunneonigrum, Apion 

bryoniae, Liriomyza 

Bucculatrix sp. Lep.: 

Bucculatricidae 39,49 

buniae phaloe, Ascia

Caicella calchas, see Cogia 

calchas 

calamistis, Sesamia 

calchas, Caicella 

calchas, Cogia 

Calephelis Lep.: Lycaenidae 

162 

Calephelis laverna Godman 

and Salvin Lep.: 

Lycaenidae 40 

Caliothrips ipomoeae 

Moulton Thy.: Thripidae 

14 

Calycomyza Dip.: 


Calycomyza allecta 

(Melander) Dip.: 

Agromyzidae 29,3 1 

Calycomyza mikaniae 

Spencer Dip.: 

Agromyzidae 128 

Calycomyza platyptera 

(Thornson) Dip.: 


campanulicollis, Promecops 

Capitophorus hippophaes 

(Walker) Hem.: Aphididae 

13 

carduana,Lobesia 

carduana, Polychrosis 

carduellinus, Hyperomyzus 

cariniventris, Rhodobaenus 

Cannenta mimosa Eichlin 

and Passoa Lep.: Sesiidae 

147,150,152-154,156 

Caryedes pickeli (Pic) Col.: 

Bruchidae 101 

Cassida exilis Col.: 

Chrysomelidae 22 

Cassida nigriventris Col.: 


Catantops humilis Serville 

Ort.: Acrididae 133 

catenulata, Lophocampa 

caulophaga, Liriomyza 

Cecidochares fluminensis 

(Lima) Dip.: Tephritidae 

40 

Cecidomyia Dip.: 

Cecidomyiidae 193 

6 Index of scientific names of insects 279 

Cecidomyia penniseti Felt 

Dip.: Cecidomyiidae 193, 

194 

Centrotypus flexuosus 


Membracidae 133 

Centrinaspis Col.: 

Curculionidae 29,30,40 

Centrinaspis perscitus see 

Linogeraeus perscitus 

Ceresa ustulata Fairmaire 

Hem.: Membracidae 143 

Cerodontha muscina 

(Meigen) Dip.: 


cervinus, Pantomorus 

Ceutorhynchus Col.: 


Ceutorhynchus asperulus 

Faust Col.: Curculionidae 

22 

Ceutorhynchus oleracae 

Marshall Col.: 

Curculionidae 2 1 3 

Ceutorhynchus portulacae 

Marshall Col.: 

Curculionidae 2 1 1,213 

Chaetogeoica graminiphaga 

Raychaudhuri, Pal and 

Ghosh Hem.: Aphididae 

0 0 

00 

Chalcodermus Col.: 

Curculionidae 144, 162 

Chalcodermus segnis Fielder 


Chalcodermus serripes 

Fahraeus Col.: 

Curculionidae 15 1, 154 

Chalcophana viridipennis 

Germar Col.: 


Chariesterus armatus 

(Thunberg) Hem.: 

Coreidae 101 

charitonia, Heliconius 

Chilo Lep.: Pyralidae 225 

Chilo agamemnon Bleszynski 

Lep.: Pyralidae 65 

Chilo zacconius Bleszynski 


Chlamisus Col.: 

Chrysomelidae 16 1 

Chlamisus insularis (Jacoby) 

Col.: Chrysomelidae 49 

Chlamisus mimosae Karren 

Col.: Chrysomelidae 30, 

39,49,150,152,154 

Chloropteryx Lep.: 

Geometridae 129 

Chnootriba similis 

(Thunberg) Col.: 

Coccinellidae 225 

Chortogonus trachypterus 

Blanchard Ort.: Acrididae 

105 

Chromatomyia Dip. 


Chrysolina Col.: 


Cicadulina triangula Hem.: 


cinerea, Piesma 

Circulifer haematoceps 

(Mulsant and Rey) Hem.: 

Cicadellidae 21 1 

citri, Planococcus 

citricola, Aphis 

civiloides, Exorista 

Cladochaeta nebulosa 

Coquillett Dip.: 

Drosophilidae 3 1 

claripennis, Phorocera 

claudina, Tegosa 

Cleitodiplosis graminis 

(Tavares) Dip.: 

Cecidomyiidae 1 8 1, 182 

Cletusfuscescens Walker 

Hem.: Coreidae 21 

Clinodiplosis Dip.: 

Cecidomyiidae 39, 40,49 

Clinodiplosis eupatorii (Felt) 


Clovia conifer (Walker) 

Hem.: Cercopidae 133 


(GuenCe) Lep.: Pyralidae 

65,90,176,182,194 

Cnaphalocrocis patnalis 

Bradley Lep.: Pyralidae 

65,90,176,182 

Coccus Hem.: Coccidae 100

280 Biological Control of Weeds: Southeast Asian Pra 

Coccus longulus (Douglas) 

Hem.: Coccidae 161 

Coelocephalapion aculeatum 

(Fall) Col.: Apionidae 150, 

152,154 

Coelocephalapion pigrae 

Kissinger Col.: Apionidae 

151,154 


spretissimum (Sharp) Col.: 

Apionidae 154 

Coelophora bissellata 

Mulsant Col.: 

Coccinellidae 134 

Coelosternus notaticeps, see 


coffearia, Homona 

Cogia calchas (Herrich- 

Schaffer) Lep.: 

Hesperiidae 145, 162 

Colaspis Col.: Chrysomelidae 

144,181 

Colaspis aerea Lefevre Col.: 

Chysomelidae 182 

Coleophora versurella Zeller 

Lep.: Coleophoridae 21,22 

commelinae, Liriomyza 

complicata, Exema 

compositae, Uroleucon 

concinnus, Euscyrtus 

conifer, Clovia 

conjuncta, Desmogramma 

connexa, Eriopis 

connexa, Procecidochares 

consimilis, Phaedon 

Contarinia Dip.: 

Cecidomyiidae 40,88,93, 

194 

Contarinia panici (Plotnikov) 


177 

Contarinia sorghicola 

(Coquillett) Dipt.: 

Cecidomyiidae 93, 193, 

194 

corbulae, Asphondylia 

cordoi, Apanteles 

coreopsidis, Aphis 

Corythuca pellucidu Drake 

and Hambleton Hem.: 

Tingidae 101 

Corythuca socia Monte 

Hem.: Tingidae 101 

costa-limai, Bemisia 

Courteia graminis, see 

Orseolia graminis 

craccivora, Aphis 

creatonoti, Apanteles 

Creatonotos gangis 

(Linnaeus) Lep.: Arctiidae 

90,111,182 

crenulata, Atractomorpha 

cretatus, Onychylis 

cretatus, Pistiacola 

cretica, Sesamia 

cribarius, Euschistus 

Criotettrix Ort.: Tetrigidae 

200,202 

Cropia minthe (Dyar) Lep.: 

Noctuidae 3 1 

cruttwellae, Neolasioptera 

Cryptocephalus miserabilis 

Suffrian Col.: 


Cryptocephalus viridiaeneus 

Boheman Col.: 


Cryptorhynchus Col.: 


cubana, Heteropsylla 

curvifascia, Erastroides 

cyanea, Altica 

cyanea, Haltica 

Cyanotricha necyria (Felder 

and Rogenhofer) Lep.: 

Dioptidae 188 

cydno, Heliconius 

cymoides, Nysius 

cynodontis. Orseolia 

Cyrtorhinus lividipennis 

Reuter Hem.: Miridae 64 

Dactylispa bipartista Col.: 


Dactynotus ambrosiae, see 

Uroleucon ambrosiae 

Dactynotus, see Uroleucon 

Delphacodes idonea Hem.: 

Delphacidae 176 

densa, Bellura 

depunctalis, Nymphula 

Desmogramma 130 

Desmogramma bigaria 

Erichson Col.: 


Desmogramma conjuncta 

BechynC Col.: 

Chrysomelidae 127, 130 

Diacamma rugosum (Le 

Guillou) Hym.: Formicidae 

43 

Diachrysia orichalcea, see 

Thysanoplusia orichalcea 

Dialeurodes vulgaris Singh 

Hem.: Aleyrodidae 30 

Dichelops fircatus 


Pentatomidae 144 

Dichomeris Lep.: Gelechiidae 

14,40 

Dicladispa armigera 

(Olivier) Col.: 


Didonis biblis (Fabricius) 

Lep.: Nymphalidae 101 

diemenalis, Lamprosema 

dilatatum, Sipylus 

diminutalis, Nymphula 

diminutalis, Paraponyx 

Dinarmus Hym.: 

Pteromalidae 152 

Dione juno (Cramer) Lep.: 

Nymphalidae 186, 1 87 

Dioxyna picciola, see 

Dioxyna sororcula 

Dioxyna sororcula 

(Wiedemann) Dip.: 

Tephritidae 3 1 

Diplacaspis miserabilis, see 

Cryptocephalus 

miserabilis 

Diplacaspis prosternalis 

(Schaeffer) Col.: 


Disonycha argentiniensis 

BechynC Col.: 


distincta, Lamprosema 

dolosa, Pegomya 

dorsalis, Bactrocera 

dorsalis, Recilia 

drumalis, Petrophila 

Dryas julia (Fabricius) Lep.: 

Nymphalidae 187

Dynamine artemisia 

(Fabricius) Lep.: 

Nymphalidae 101 

Dyodiplosis andropogonis, 

see Orseolia andropogonis 

Dyodiplosisfluitans, see 

Lasioptera fluitans 

Dyodiplosisfluvialis, see 

Orseolia fluvialis 

Dyops minthe, see Cropia 

minthe 

echeclus, Hippotion 

echinochloa, Lasioptera 

Echinocnemus Col.: 

Curculionidae 1 15 

Echoma marginata 

(Linnaeus) Col.: 

Chrysomelidae 127, 1 30 

Echoma quadristillata 

(Boheman) Col.: 


Ecpantheria hambletoni, see 

Hypercompe hambletoni 

Edessa meditabunda 



eichhorniae, Neochetina 

Elachertus Hym.: Eulophidae 

46 

Elasmopalpus lignosellus 

(Zeller) Lep.: Pyralidae 65 

Elasmus Hym.: Elasmidae 46 

Eldana saccharins Walker 


elegans, Neohydronomus 

elimatus, Dalbulus 


(Walker) Lep.: Pyralidae 

115,116,167,201,202 

ementitus, Xyonysius 

Emmalocera Lep.: Pyralidae 

63,65 

Enchenopa gracilis (Germar) 


entreriana, Tomaspis 

entreriana, Zulia 

Entylia Hem.: Membracidae 

128 

Entylia sinuata (Fabricius) 



Epilachna indica Mulsant 

Col.: Coccinellidae 134 

Epilachna similis, see 

Chnootriba similis 

Epipagis albiguttalis, see 

Sameodes albiguttalis 

Epipsammia pectinicornis, 

see Spodoptera 

pectinicornis 

Episammea pectinicornis, see 

Epipsammia pectinicornis 

Episcada pascua Schaus 


Erastroides curvifascia 

Hampson Lep.: Noctuidae 

200 

erato, Heliconius 

Eretmocera impactella 

(Walker) Lep.: 

Scythrididae 23 

eridania, Prodenia 

eridania, Spodoptera 

Erinnyis oenotrus (Cramer) 

Lep.: Sphingidae 101 

Eriopis connexa (Germar) 

Col.: Coccinellidae 141 

Eryphanis polyxena 

Meerburgh Lep.: 

Nymphalidae 193, 194 

erythrocephalus, Stiretrus 

etheiella, Eucampyla 

Eublemma versicolor, see 

Autoba versicolor 

Eucalymnatus Hem.: 

Coccidae 100 

Eucampyla etheiella Meyrick 

Lep.: Pyralidae 46,47 

Euderus Hym.: Eulophidae 

52 

Eueides aliphera Godart 


Eueides isabella (Cramer) 


Eueides procula Butler and 

Drull Lep.: Nymphalidae 

187 

eupatoriella, Melanagromyza 

eupatorii, Clinodiplosis 

eupatorii, Neolasioptera 

euphorbiae, Haplothrips 

Eupithecia Lep.: Geometridae 

129 

Eurema Lep.: Pieridae 139 

Eurema lisa (Boisduval and 

LeConte) Lep.: Pieridae 

162 

Eurema tenella (Boisduval) 

Lep.: Pieridae 145, 162 

Euschistus tristigmus 

cribarius Stdl Hem.: 


Euschistus luridus Dallas 

Hem.: Pentatomidae 144 

Euscyrtus concinnus (de 

Haan) Ort.: Gryllidae 57, 

64,224 

Exema complicata Jacoby 


132 

exigua, Spodoptera 

exilis, Cassida 

Exorista Dip.: Tachinidae 43 

Exorista civiloides, see 

Exorista xanthaspis 

Exorista xanthaspis 

Wiedemam Dip.: 

Tachinidae 45 

explanata, Pentispa 

fabae, Aphis 

fasciatus, Pistiacola 

fascifrons, Macrosteles 

fastigiata, Sibinia 

fatualis, Ategumia 

femorata, Sagra 

Ferrisia virgata (Cockerell) 

Hem.: Pseudococcidae 105 

ferrugenea, Bothrogonia 

ferruginae, Temlucha 

flexuosus, Centrotypus 

floris, Melanagromyza 

fluitans, Dyodiplosis 

fluitans, Lasioptera 

fluitantis, Apanteles 

fluminensis, Cecidochares 

fluvialis, Dyodiplosis 

fluviatilis, Orseolia 

fregonalis, Nymphula 

frugiperda, Spodoptera 

frugivora, Neolasioptera 

furcatus, Dichelops


furcifera, Sogatella 

furnacalis, Ostrinia 

fuscesens, Cletus 

gangis, Amsacta 

gangis, Creatonotos 

ganglbaueri, Haplothrips 

Garcanus gracilentus (St&) 

Hem.: Miridae 30 

Gargara Hem.: Membracidae 

121 

geminata, Solenopsis 

gentilei, Tetrastichus 

gentilei, Thripsastichus 

Geoica Iucifuga (Zehntner) 


Germalus unipunctatus 

(Montandon) Hem.: 

Lygaeidae 21 

Geromyia penniseti (Felt) 


194 

Geromyia seminis (Felt) Dip.: 

Cecidomyiidae 193,194 

Gesonula punctifrons (Sdl) 

Ort.: Acrididae 78, 167 

Gibbobruchus pickeli , see 

Caryedes pickeli 

Gibbobruchus polycoccus 

(Fahraeus) Col.: Bruchidae 

101 

godmani, Asynonychus 

gossypii, Aphis 

gowdeyi, Haplothrips 

gracilentus, Garcanus 

gracilis, Enchenopa 

graminiphaga, Chaetogeoica 

graminis, Cleitodiplosis 

graminis, Courteia 

graminis, Orseolia 

graminum, Schizaphis 

granarium, Macrosiphum 

griseola, Hydrellia 

griseus, Brachycyttarus 

guerini, Lexiphanes 

guildinii, Piezodorus 

gunniella, Neurostrota 

Gyrocnemis Hem.: 

Thyreocoridae 144 

haematoceps, Circulijier 

I 

Haimbachia infusella 

(Walker) Lep.: Pyralidae 

69,73,74,76,79,80,82 

Haltica cyanea, see Altica 

cyanea 

Halticus minutus Reuter 


hambletoni, Ecpantheria 

hambletoni, Hypercompe 

hanno, Hemiargus 

Haplopeodes Dip.: 

Agromyzidae 21 6 

Haplopeodes minutus (Frost) 

Dip.: Agromyzidae 21,22 


(Czoquillett) Dip.: 

Agromyzidae 2 12,2 1 3, 

216 

Haplothrips Thy.: 

Phlaeothripidae 134 

Haplothrips euphorbiae Thy.: 


Haplothrips ganglbaueri 

Schmutz Thy.: 

Phlaeothripidae 63, 64, 89 

Haplothrips gowdeyi 

(Franklin) Thy.: 

Phlaeothripidae 14, 101, 

182 

Haplothrips longisetosus 

Ananthakrishnan Thy.: 


Haplothrips paumalui 

Moulton Thy.: 


harleyi, Teleonemia 

hawaiiensis, Thrips 

hecale, Heliconius 

helichrysi, Brachycaudus 

Heliconius charitonia 

(Linnaeus) Lep.: 


Heliconius cydno Bates Lep.: 


Heliconius erato Doubleday 


Heliconius hecale Hewitson 

Lep.: Nymphalidae 187, 

189 

Helicoverpa obsoleta, see 

Helicoverpa zea 

Helicoverpa virescens, see 

Heliothis virescens 

Helicoverpa zea (Boddie) 


Heliodines Lep.: Heliodinidae 

216 

Heliodines quinqueguttata 

Walshingham Lep.: 

Heliodinidae 2 13,2 16 

Heliothis virescens 


Noctuidae188 

Hellula undalis (Fabricius) 


Helopeltis Hem.: Miridae 133 

Helopeltis antonii Signoret 


Hemiargus hanno (Stoll) 

Lep.: Lycaenidae 145, 161, 

162 

Hemiberlesia lataniae 

(Signoret) Hem.: 

Diaspididae 161 

hennia, Proxenus 

herbidum, Acrosternum 

Heteropsylla Hem.: Psyllidae 

139,142,144,161 

Heteropsylla cubana 

Craw ford Hem.: Psyllidae 

1 42 

Heteropsylla spinulosa 

Muddiman, Hodkinson and 

Hollis Hem.: Psyllidae 

137,139-1 45 

hieroglyphicus, Poekilocerus 

Hilar Col.: Chrysomelidae 

1 44 

Hippotion echeclus 

(Boisduval) Lep.: 

Sphingidae 78, 167 

hirsuta, Tetraneura 

hirticornis, Aerenica 

Homoeocerus serrijier 

Westwood Hem.: Coreidae 

133 

Homona cofearia (Nietner) 

Lep.: Tortricidae 45 

Horciacinus argentinus, see 

Horciacinus signoreti 

Horciacinus signoreti (Berg) 

Hem.: Miridae 143

Horcias nobilellus (Berg) 

Hem.: Miridae 22,30 

Horismenus ?aeneicollis 

Ashmead Hym.: 

Eulophidae 130, 132 

hospes, Spilostethus 

huebneri, Amata 

humilis, Catantops 

humilis, Hyperodes 

humilis, Listronotus 

Hyalomyodes triangulifer 

(Loew) Dip.: Tachinidae 

130 

Hyalopeplus vitripennis (Stll) 


Hydrellia griseola (Falltn) 

Dip.: Ephydridae 65 

Hymenia recurvalis, see 

Spodoptera recurvalis 

Hypanthus Col.: 


Hypercompe hambletoni 

(Schaus) Lep.: Arctiidae 

3 1 

Hyperodes humilis, see 

Listronotus humilis 

Hyperomyzus carduellinus 

(Theobald) Hem.: 

Aphididae 13 

Hypolixus ritsemae (Pascoe) 


Hypolixus trunculatus 

(Fabricius) Col.: 

Curculionidae 19,21,22, 

45 

Hypomicrogaster Hym.: 

Braconidae 81 

Hypurus bertrandi Pems 

Col.: Curculionidae 

21 1-213,215-217 

Hysteroneura setariae 

(Thomas) Hem.: 

Aphididae 89 

Icerya seychellarum 

(Westwood) Hem.: 

Margarodidael61 

Idiophantis Lep.: Gelechiidae 

122 

idonea, Delphacodes 

illinoisensis, Aphis 


Imerodes Col.: Curculionidae 

122 

impactella, Eretmocera 

ina, Taractrocera 

inaequalis, Xyonysius 

includens, Pseudoplusia 

inconspicuus, Nysius 

incurvus, Pycnoderes 

indica, Epilachna 

indica, Mycodiplosis 

infusella, Acigona 

infusella, Haimbachia 

insecta, Xanthaciura 

insignis, Liriomyza 

insularis, Chlamisus 

insulata, Pareuchaetes 

insulata, Ammalo 

inustorum, Lasioptera 

ipomoeae, Caliothrips 

ipsilon, Agrotis 

irrorata. Milothris 

isabella, Eueides 

Isothrips Thy.: Thripidae 1 33 

Itonida penniseti, see 

Geromyia penniseti 

Itonida seminis, see 

Geromyia seminis 

janata, Achaea 

javanica, Parallelodiplosis 

julia, Dryas 

juno, Dione 

kanni, Lasioptera 

knysna, Zizeeria 

kolophon, Sogatella 

krupta, Zizeeria 

Lactica sp. Col.: 


lactinea, Amsacta 

laevis, Macrotracheliella 

Lamprosema diemenalis 

(Guenke) Lep.: Pyralidae 

132,133 

Lamprosema distincta (Kaye) 


Laodelphax striatellus 

(Falltn) Hem.: 

Delphacidae 64,89,93 

laphygmae, Meteorus 

Lasioptera Dip.: 

Cecidomyiidae 18 1 

Lasioptera echinochloa Felt 


65 

Lasiopterafluitans Felt Dip.: 


Lasioptera inustorum Felt 


Lasioptera kanni Felt Dip.: 


Lasioptera panici Felt Dip.: 


Lasioptera paniculi Felt Dip.: 


Lasioptera portulaceae Felt 

Dip.: Cecidomyiidae 21 3 

lataniae, Hemiberlesia 

latigenis, Notiphila 

latipes, Mocis 

latiuscula, Scotinophara 

laverna, Calepheles 

leelamaniae, Macrosiphum 

leelamaniae, Sitobion 

Leptocentrus taurus 


Membracidae 12 1 

Leptocorisa acuta (Thunberg) 

Hem.: Alydidae 99 

Leptocorisa oratorius 


Alydidae 64,99 

Leptocorisa solomonensis 

Ahmad Hem.: Alydidae 99 

Leptocysta sexnebulosa (Stll) 


leucaenae, Tamarixia 

leucopterus, Blissus 

leuculias, Argyresthia 

Lexiphanes Col.: 


Lexiphanes guerini (Perbose) 

Col.: Chrysomelidae 15 1 

Lexiphanes semicyaneus 

(Suffrian) Col.: 


licarsisalis, Herpetogramma 

Eignosellus, Elasmopalpus 

linearis, Riptortus 

lineatus, Astylus 

Linogeraeus perscitus 

(Herbst) Curculionidae 2 13


Liothrips mikaniae (Priesner) 

Thy.: Phlaeothripidae 125, 

127,130,134,135 

liothrips, Thripsobremia 

Liriomyza Dip.: Agromyzidae 

31,32 

Liriomyza archboldi Frost 

Dip.: Agromyzidae 29,3 1 

Liriomyza bryoniae 

(Kaltenbach) Dip.: 


Liriomyza caulophaga 

(Kleinschmidt) Dip.: 

Agromyzidae 21 1,212 

Liriomyza commelinae Frost 


Liriomyza insignis Spencer 

Dip.: Agromyzidae 29,3 1 

Liriomyza marginalis 

Malloch Dip.: 


Liriomyza robustae Spencer 


Liriomyza strigata Meigen 


Liriomyza trifolii (Burgess) 

Dip.: Agromyzidae 3 1,211 

Liriomyza venegasiae 

Spencer Dip.: 


lisa, Eurema 

Lissorhoptrus oryzophilus 

Kuschel Col.: 


Listronotus humilis 

(Gyllenhall) Col.: 


litura, Spodoptera 

lividipennis, Cyrtorhinus 

Lixophaga Dip.: Tachinidae 

206 

Lobesia carduana (Busck) 


longifircifera, Sogatella 

longisetosus, Haplothrips 

Longitarsus Col.: 

Chrysomelidae 13 1, 132 

Longitarsus nr amazonus 

Baly Col.: Chrysomelidae 

128,132 

longulus, Coccus 

Lophocampa catenulata 

(Hubner) Lep.: Arctiidae 

161,162 

lorata, Baris 

Loxostege Lep.: Pyralidae 23 

Loxostege bijidalis 

(Fabricius) Lep.: Pyralidae 

212 

lucifuga, Geoica 

lugens, Nilaparvata 

luridus, Euschistus 

luteirostre, Apion 

macminni, Cerodontha 

rnacrophthalma, Diopsis 

Macrosiphum granarium, see 

Sitobion avenue 

Macrosiphum leelamaniae, 

see Sitobion leelamaniae 

Macrosiphum miscanthi, see 

Sitobion miscanthi 

Macrosiphum solidaginis, see 

Uroleucon solidaginis 

Macrosteles fascifrons (Still) 

Hem.: Cicadellidae 167 

maculosa, Amauromyza 

Maecolaspis aerea, see 

Colaspis aerea 

maidis, Dalbulus 

maidis, Peregrinus 

malayanus, Nephotettix 

Mansonia Dip.: Culicidae 

199 

Marasmia patnalis, see 

Cnaphalocrocis patnalis 

marginalis, Liriomyza 

marginata, Echoma 

Masonaphis anaphalidis, see 

Neomasonavhis 

anaphalidis 

mauritia, Spodoptera 

mazans, Staphylus 

medinalis, Cnaphalocrocis 

meditabunda, Edessa 

Melanagromyza Dip.: 


Melanagromyza bidentis 

Spencer Dip.: 


Melanagromyza eupatoriella 

Spencer Dip.: 

Agromyzidae 39,44,45 

Melanagromyza floris 

Spencer Dip.: 


Melanagromyza metallica 

(Thomson) Dip.: 

Agromyzidae 13, 14 

Melanagromyza polyphyta 

Kleinschmidt Dip.: 

Agromyzidae 1 88 

Melanagromyza splendida 

Frick Dip.: Agromyzidae 

3 1 

rneridionalis, Arthrocnoda. 

Mescinia parvula (Zeller) 

Lep.: Pyralidae 39,42,44, 

46,49,50 

metallica, Melanagromyza 

Metqonycha pallidula 

(Boheman) Col.: 


Meteorus Hym.: Braconidae 

121 

micaceana, Archips 

Microcephalothrips Thy.: 

Thripidae 133 


abdominalis (D.L. 

Crawford) Thy.: Thripidae 

14 

Microcomps~s, see 

Microporus 

Microporus Hem.: Cydnidae 

161 

Micrutalis Hem.: 


Micrutalis binaria 

(Fairmaire) Hem.: 


mikaniae, Calycomyza 

mikaniae, Liothrips 

Milothris irrorata (Fabricius) 

Col.: Cerambycidae 153 

mimosa, Carmenta 

mimosae, Chlamisus 

minimus, Argentinorhynchus 

minthe, Cropia 

minthe, Dyops 

minuta, Oxya 

minutispina, Nilautama 

minutus, Halticus 

minutus, Haplopeodes

misantlensis, Pareuchaetes 

miscanthi, Macrosiphum 

miscanthi, Sitobion 

miserabilis, Cryptocephalus 

miserabilis, Diplacaspis 

Mocis latipes (GuenCe) Lep.: 

Noctuidae 3 1 

Monecphora bicincta 

fraterna, see Prosapia 

bicincta fraterna 

multiplicalis, Samea 

multistrigata, Phalonidia 

muscina, Cerodontha 

Mycetaspis personata 

(Comstock) Hem.: 

Diaspididae 14 

Mycodiplosis indica Felt 


Myrmicaria brunnea 

Saunders Hym.: 

Formicidae 43 

Mythimna unipuncta 

(Haworth) Lep.: Noctuidae 

65 

Myzus ornatus Laing Hem.: 

Aphididae 13 

Myzus persicae (Sulzer) 

Hem.: Aphididae 13,21, 

24, 188., 21 1,224 

Namangana pectinicornis, 

see Epipsammia pectini 

cornis 

nebulosa, Cladochaeta 

necyria, Cyanotricha 

Neochetina Col.: 

Curculionidae 73,76,78 

Neochetina bruchi Hustache 

Col.: Curculionidae 69,73, 

74,76-82 

Neochetina eichhorniae 

Warner Col.: 

Curculionidae 69,73-80, 

82 

Neogalea sunia (GuenCe) 

Lep.: Noctuidae 22,212 

Neohydronomus afJinis 

Hustache Col.: 

Curculionidae 197, 199, 

200,202-206 


Neohydronomus elegans 



Neohydronomus pulchellus 



Neolasioptera Dip.: 

Cecidomyiidae 128, 132, 

217 

Neolasioptera brickelliae 


40 

Neolasioptera cruttwellae 

Gagne Dip.: 


Neolasioptera eupatorii (Felt) 


Neolasioptera frugivora 

Gagne Dip.: 


Neolasioptera portulacae 

(Cook) Dip.: 

Cecidomyiidae 21 3,2 17 

Neomasonaphis anaphalidis 

(Basu) Hem.: Aphididae 

13 

Nephotettix Hem.: 


Nephotettix malayanus 

Ishihara and Kawase 


Nephotettix nigropictus, see 

Nephotettix 

nigromaculatus 

Nephotettix nigromaculatus 

(Motschulsky) Hem.: 

Cicadellidae 89,225 

Nephotettix virescens 

(Distant) Hem.: 


nervosa, Nisia 

Nesoclutha pallida (Evans) 


Neurostrota gunniella 

(Busck) Lep.: 

Gracillariidae 147, 150, 

152,154156,162 

nigriabdominalis, Tetraneura 

nigricauda, Aphis 

nigrifrons, Graminella 

nigrirostris, Pistiacola 

nigriventris, Cassida 

nigromaculatus, Nephotettix 

nigropictus, Nephotettix 

Nilaparvata lugens (StAl) 

Hem.: Delphacidae 64 

Nilautama minutispina 

Funkhouser Hem.: 


Nisaga simplex Walker Lep.: 

Eupterotidae 176 

Nisia atrovenosa, see Nisia 

nervosa 

Nisia nervosa (Motschulsky) 

Hem.: Meenoplidae 199, 

200,202 

Nisoniades bessus, see Cogia 

calchas 

noacki, Platinglisia 

nobilellus, Horacias 

Nodonota Col.: 


notaticeps, Coelosternus 

notaticeps, Sternocoelus 

Notiphila latigenis Hendel 

Dip.: Ephydridae 1 15,116 

Notophila similis de Meijere 

Dip.: Ephydridae 1 15,116 

nymphaeae, Rhopalosiphum 

Nymphula depunctalis, see 


Nymphula diminutalis 

Snellen, see Parapoynx 

diminutalis 

Nymphulafregonalis Snellen 


Nymphula responsalis, see 


Nymphula tenebralis, see 

Parapoynx tenebralis 

Nymphula turbata, see 

Parapoynx turbata 

Nysius Hem.:Lygaeidae 21 

Nysius cymoides (Spinola) 

Hem.: Lygaeidae 21 1 

Nysius inconspicuus Distant 

Hem.: Lygaeidae 14, 105 

Nysius vinitor Bergroth 

Hem.: Lygaeidae 21 1 

obliteralis, Synclita 

obsoleta, Helicoverpa

286 


Ochetina bruchi Hustache 


ochreosa, Sibinia 

Odontomachus simillimus Fr. 

Smith Hym.: Formicidae 

43 

Oecophylla smaragdina 

(Fabricius) Hym.: 

Formicidae 43 

oenotrus, Erinnyis 

Olcella pleuralis Becker 

Dip.: Chloropidae 14 

oleracae, Ceutorhynchus 

Omoplata rnarginata, see 

Echoma marginata 

Onebala tegulella 

(Walshingham) Lep.: 

Gelechiidae 128 

. Onychylis cretatus, see 


Onychylis sp. nr nigrirostris, 

see Pistiacola sp. nr 

nigrirostris 

oo, Plusia 

oratorius, Leptocorisa 

orichalcea, Diachrysia 

orientalis, Orosius 

ornatus, Myzus 

Orosius albicinctus, see 



(Matsumura) Hem.: 

Cicadellidae 21 1 

Orseolia Dip.: Cecidomyiidae 

87 

Orseolia andropogonis Felt 


Orseolia cynohntis Kieffer 

and Massalongo Dip.: 


Orseoliafluvialis Dip.: 


Orseoliafluviatilis (Felt) 


90 

Orseolia graminis (Kreffer 

and Docters Van Leeuwen- 

Reijnvaan) Dip.: 


Orseolia oryzae (Wood- 

Mason) Dipt.: 

Cecidomyiidae 65,90, 

175,177 

oryzae, Orseolia 

oryzae, Pachydiplosis 

oryzivora, Thaia 

oryzophilus, Lissorhoptrus 

Ostriniafurnacalis (GuenCe) 


ovata, Brachymeria 

Oxya minuta Carl Ort.: 

Acrididae 78 

Oxysychus Hym.: 

Pteromalidae 21 

Pachybrachys Col.: 


Pachydiplosis, see Orseolia 

SP- 

Pachydiplosis oryzae, see 

Orseolia oryzae 

pacificus, Brachyplatys 

pacificus, Planococcus 

palegon, Thecla 

palegon, Thereus 

palliata, Phytomyza 

palliatus, Haplopeodes 

pallida, Nesoclutha 

pallidula, Metaxyonycha 

pallidulus, Taylorilygus 

panici, Contarinia 

panici, Lasioptera 

panici, Stenodiplosis 

panicola, Sogatella 

paniculi, Lasioptera 

Pantomorus cervinus, see 

Asynonychus godrnani 

Parallelodiplosis Dip.: 


Parallelodiplosis javanica 

Felt Dip.: Cecidomyiidae 

177 

Parallelodiplosis paspali 

Dip.: Cecidomyiidae 18 1 

Parapoynx diminutalis 

Snellen Lep.: Pyralidae 

20 1 

Parapoynx stagnalis (Zeller) 

Lep.: Pyralidae 176, 182 

Parapoynx tenebralis 

(Lower) Lep.: Pyralidae 

199,20 1 

Parapoynx turbata Butler 

Lep.: Pyralidae 199,201 

Pareuchaetes Lep.: Arctiidae 

38,40 

Pareuchaetes arravaca 

(Jordan) Lep.: Arctiidae 40 

Pareuchaetes aurata 

aurantior Rothschild Lep.: 

Arctiidae 40 

Pareuchaetes aurata aurata 

(Butler) Lep.: Arctiidae 40, 

42,47,50 


(Walker) Lep.: Arctiidae 

40 

Pareuchaetes misantlensis 

Rego Barros Lep.: 

Arctiidae 40 

Pareuchaetes pseuhinsulata 

Rego Barros Lep.: 

Arctiidae 14,35,38,39, 

42-47,50,52 

Parnara bada (Moore) Lep.: 

Hesperiidae 194 

Parthenothrips Thy.: 

Thripidae 134 

parvicornis, Agromyza 

parvula, Mescinia 

parvus, Phenacoccus 

pascua, Episcada 

paspali, Parallelodiplosis 

patnalis, Cnaphalocrocis 

patnalis, Marasmia 

Paulinia acuminata De Geer 

Ort.: Acrididae 200 

paumalui, Haplothrips 

pectinicornis, Epipsammia 

pectinicornis, Spohptera 

pedestris, Bactrocera 

Pegomya dolosa Stein Dip.: 

Anthomyiidae 21 3,217 

pellucida, Corythuca 

pennatula, Psalis 

penniseti, Cecidomyia 

penniseti, Geromyia 

penniseti, ltonida 

penniseti, Sesamia 

Pentispa explanata (Chapuis) 


5 1 

Perasphondylia reticulata 


39,51

perditor, Thyanta 

Peregrinus maidis (Ashmead) 

Hem.: Delphacidae 89,225 

perlata, Ptychamalia 

Perrhybris phaloe (Godart) 

Lep.: Pieridae 3 1 

perscitus, Centrinaspis 

perscitus, Linogeraeus 

persicae, Myzus 

personata, Micetaspis 

pertinax, Phaedon 

peruana, Sibinia 

Petrophila drumalis (Dyar) 


Phaedon consimilis, see 

Phaedon pertinax 

Phaedon pertinax StAl Col.: 


phaloe, Perrhybris 

Phalonidia multistrigata 

Walshingham Lep.: 

Tortricidae 129 

philippinensis, 

Pseudonapomyza 

Pholetesor sp. 

(circumscriptus group) 

Hym.: Braconidae 2 1 6 

Phorocera claripennis Pales 

Dip.: Tachinidae 187 

Physimerus Col.: 


Physimerus pygmaeus Jacoby 


127,131 

Phytomyza atricornis Meigen 

Dip.: Agromyzidae 3 1 

Phytomyza palliata, see 


picciola, Dioxyna 

pickeli, Caryedes 

pickeli, Gibbobruchus 

pieridis, Vesiculaphis 

Piesma cinereum Say Hem.: 

Piesmatidae 22 

Piezodorus guildinii 

(Westwood) Hem.: 


pigrae, Coelocephalapion 

pigricola, Acanthoscelides 

pilicornis, Schizocerella 


Pinnaspis strachani (Cooley) 

Hem.: Diaspididae 16 1 

Pionea upalusalis (Walker) 



(Champion) Col.: 


Pistiacola fasciatus Wibmer 

and O'Brien Col.: 


Pistiacola sp. nr nigrirostris 


Planococcus citri (Risso) 

Hem.: Pseudococcidae 200 

Platinglisia noacki Cockerel1 

Hem.: Coccidae 101 

Platynota rostrana (Walker) 


platyptera, Calycomyza 

pleuralis, Olcella 

Plusia oo , see Pseudoplusia 

includens 14 

Plutella xylostella (Curtis) 

Lep.: Yponomeutidae 23 

Pochazia antica (Gray) 

Hem.: Ricaniidae 121 

Podogaster Hym.: 


Poekilocerus hieroglyphicus 

(Klug) Ort.: Acrididae 99 

Polychrosis ?carduana, see 

hbesia ?carduana 

polycoccus, Gibbobruchus 

polyphyta, Melangromyza 

polyxena, Eryphanis 

portulacae, Asphondylia 

portulacae, Baris 

portulacae, Ceutorhynchus 

portulacae, Neolasioptera 

portulaecae, Lasioptera 

Procecidochares Dip.: 

Tephritidae 39 

Procecidochares connexa 

Macquart Dip.: Tephritidae 

40,44,51 

procula, Eueides 

Prodenia eridania, see 


prolixa, Teleonemia 

Promecops Col.: 

Curculionidae 29,30, 144 

Promecops campanulicollis 

Voss Col.: Curculionidae 

162 

Prosapia bicincta fraterna 

(Uhler) Hem.: Cercopidae 

89 

proserpina, Tarophagus 

prosternalis, Diplacaspis 

Protonectarina sylveiriae 

(Saussure) Hym.: Vespidae 

141 

Proxenus hennia Swinhoe 


proxirna, Agromyza 

Psalis pennatula (Fabricius) 

Lep.: Lymantriidae 225 

Pseudaletia unipuncta, see 

Mythimna unipuncta 

Pseudoderelomus 

baridiiformis Champion 

Col.: Curculionidae 127, 

131 

pseudoinsulata, Pareuchaetes 

pseudoinsulata, Pareuchaetes 

Pseudonapomyza 

philippinensis Spencer 


Pseudonapomyza spicata 

Malloch Dip.: 


Pseudoplusia includens 

(Walker) Lep.: Noctuidae 

1 " 

14 

Psigida walkeri (Grote) Lep.: 

Cercophanidae 137, 139, 

141,144 

Psyllaephagus yaseeni Noyes 

Hym.: Encyrtidae 141, 142 

Psylopigida walkeri, see 

Psigida walkeri 

Pteronemobius Ort.: 

Gryllidae 224 

Ptychamalia perlata 

(Warren) Lep.: 

Geometridae 162 

pulchellus, Neohydronomus 

punctifrons, Gesonula 

puniceus, Acanthoscelides 

pusanus, Sogatodes 

pusanus, Tagosodes

288 


Pycnoderes incurvus 

(Distant) Hem.: Miridae 

128 

pygmaeus, Physimerus 

quadridentatus, 


quadristillata, Echoma 

querci, Anoecia 

querci, Stegophylla 

quinqueguttata, Heliodines 

Rastrococcus Hem.: 

Pseudococcidae 121 

Recilia dorsalis 

(Motschulsky) Hem.: 


, recurvalis, Hymenia 

recurvalis, Spodoptera 

recurvalis, Spoladea 

Recurvaria Lep.: Gelechiidae 

128 

responsalis, Elophila 

responsalis, Nymphula 

reticulata, Peraphondylia 

Rhodobaenus cariniventris 

Champ. Col.: 

Curculionidae 30,39,47, 

51,52 

Rhodobaenus 

tredecimpunctatus (Illiger) 


Rhopalosiphum nymphaeae 

(Linnaeus) Hem.: 

Aphididae 77,200,201 

Riptortus linearis (Linnaeus) 

Hem.: Coreidae 133 

ritsemae, Hypolixus 

robustae, Liriomyza 

rostrana, Platynota 

rugosum, Diacamma 

russelli, Brachymeria 

s-littera, Systena 

saccharina, Eldana 

Sagra femorata (Drury) Col.: 


Samea multiplicalis (Guente) 


201-203,205 

Sameodes Lep.: Pyralidae 76 

Sameodes albiguttalis 

(Warren) Lep.: Pyralidae 

69,73,75-82 

Scamurius Hem.: Coreidae 

137,139,140,142,143, 

145 

Sceloenopla Col.: 


Schizaphis graminum 

(Rondani) Hem.: 

Aphididae 89 

Schizocerella pilicornis 

(Holmgren) Hym.: 

Tenthredinidae 21 2,213, 

217,218 

Scotinophara latiuscula 

(Breddin) Hem.: 

Pentatomidae 57,64, 1 11 

scrupulosa, Teleonemia 

segnis, Chalcodermus 

Selca brunella (Hampson) 

Lep.: Noctuidae 120- 123 

semicyaneus, Lexyphanes 

seminicola, Sibinia 

seminis, Geromyia 

seminis, Itonida 

septentrionalis. Platysimus 

serrifer, Homoeocerus 

serripes, Chalcodermus 

Sesamia Lep.: Noctuidae 225 

Sesamia botanephaga Tams 

and Bowden Lep.: 

Noctuidae 65 

Sesamia calamistis Hampson 


Sesamia cretica Lederer Lep.: 

Noctuidae 176 

Sesamia penniseti Tams and 

Bowden Lep.: Noctuidae 

65 

setariae, Hysteroneura 

sexnebulosa, Leptocysta 

seychellarum, Icerya 

Sibinia Col.: Curculionidae 

144,153 

Sibinia aspersa Champion 


Sibinia fatigiata Clark Col.: 


Sibinia ochreosa Casey Col.: 

Curculionidae 15 1, 1 55 

Sibinia peruana Pierce Col.: 


Sibinia seminicola Clark 

Col.: Curculionidae 151, 

155 

Sibinia subulirostris 



signoreti, Horciacinus 

similis, Chnootriba 

similis, Epilachna 

similis, Notiphila 

similis, Tegosa 

simillimus, Odontomachus 

simplex, Nisaga 

sinuata, Entylia 

Sipylus Hem.: Membracidae 

121 

Sipylus dilatatum (Walker) 

Hem.: Membracidae 1 2 1 

Sitobion avenue (Fabricius) 


Sitobion leelamaniae (David) 


Sitobion miscanthi 

(Takahashi) Hem.: 

Aphididae 89 

smaragdina, Oecophylla 

soccata, Atherigona 

socia, Corythuca 

Sogatellafurcifera (Horviith) 

Hem.: Delphacidae 64, 89 

Sogatella kolophon 

(Kirkaldy) Hem.: 


Sogatella longifurcifera, see 

Sogatella vibix 

Sogatella panicicola, see 

Sogatella vibix 

Sogatella vibix (Haupt) Hem.: 


Sogatodes pusanus, see 

Tagosodes pusanus 

solani, Aulacorthum 

Solenopsis geminata 

(Fabricius) Hym.: 

Forrnicidae 1 8 1 

solidaginis, Macrosiphum 

solidaginis, Uroleucon 

solomonensis, Leptocorisa 

sorghicola, Contarinia

sororcula, Dioxyna 

Sphaeniscus atilus (Walker) 

Dip.: Tephritidae 134 

Sphenarches anisodactylus 

(Walker) Lep.: 

Pterophoridae 188, 189 

spicata, Pseudonapomyza 

Spilosoma virginica 

(Fabricius) Lep.: Arctiidae 

200,201 

spinulosa, Heteropsylla 

spiraecola, Aphis 

splendida, Melanagromyza 

Spoabptera eridania (Stoll) 

Lep.: Noctuidae 22, 101, 

212 

Spoabptera exigua (Hiibner) 

Lep.: Noctuidae 22,212 

Spodopterafrugiperda (J.E. 

Smith) Lep.: Noctuidae 14, 

90,225 

Spodoptera Iitura (Fabricius) 

Lep.: Noctuidae 22,78, 

162,167 

Spoabptera mauritia 

(Boisduval) Lep.: 

Noctuidae 78,202 

Spoabptera pectinicornis 

(Hampson) Lep.: 

Noctuidae 197,200, 

202-204,206 

Spoabptera sunia, see 

Neogalea sunia 

Spoladea recurvalis 


Pyralidae 23 

spretissimum, 


squamosus, 

Argentinorhynchus 

stagnalis, Parapoynx 

Staphylus mazans (Reakirt) 

Lep.: Hesperiidae 162 

Stenchaetothrips biformis 

Bagnall Thy.: Thripidae 64 

Stegophylla querci (Fitch) 


Stenodiplosis Dip.: 


Stenodiplosis panici, see 

Contarinia panici 


Sternocoelus Col.: 



(Marshall) Col.: 


Stiretrus erythrocephalus 

Lepeletier and Serville 

Hem.: Pentatomidae 29,30 

Stomatomyia Dip.: 

Tachinidae 1 8 1 

strachani, Pinnaspis 

striatellus, Laodelphax 

strigata, Liriomyza 

subornata, Argyractis 

subulirostris. Sibinia 

sunia, Neogalea 

sunia, Spodoptera 

Sycanus Hem.: Reduviidae 43 

sylveiriae, Protonectarina 

Synclita obliteralis (Walker) 


Syphrea bibiana Bechynk 

Jacoby Col.: 


Systena s-littera (Linnaeus) 


tabaci, Bemisia 

tabaci, Thrips 

Tagosodes pusanus (Distant) 

Hem.: Delphacidae 64 

Tamarixia Eeucaenae Boucek 

Hym.: Eulophidae 142 

Taractrocera ina Waterhouse 

Lep.: Hesperiidae 182 

Tarophagus proserpina 

(Kirkaldy) Hem.: 

Delphacidae 57, 105, 167 

taurus, Leptocentrus 

Taylorilygus pallidulus 

(Blanchard) Hem.: Miridae 

1 43 

Tegosa claudina 

(Eschscholtz) Lep.: 

Nymphalidae 129,132 

Tegosa similis, see Tegosa 

claudina 

tegulella, Onebala 

Teleonemia Hem.: Tingidae 

125,131,132 

Teleonemia harleyi 

Froeschner Hem.: Tingidae 

131 

Teleonemia prolixa St31 


Teleonemia scrupulosa Stil 


Teleonemia sp. or spp. nr 

prolixa Hem.: Tingidae 

127,131 

Temelucha ferruginae 

(Davis) Hym.: 


Temnodachrys aphodoides 


tenebralis, Nymphula 

tenebralis, Parapoynx 

tenella, Eurema 

Tetraneura basui Hille Ris 

Lambers Hem.: Aphididae 

89 

Tetraneura hirsuta, see 

Tetraneura 

nigriabdominalis 

Tetraneura nigriabdominalis 

(Sasaki) Hem.: Aphididae 

89 

Tetrastichus Hym.: 

Eulophidae 1 15, 132 

Tetrigella Hem.: Cicadellidae 

121 

Tettigoniella, see Tettigella 

Thagona tibialis (Walker) 

Lep.: Lymantriidae 101 

Thaia oryzivora Ghauri 


theae, Tropicomyza 

Thecla azia, see Tmolus azia 

Thecla palegon, see Thereus 

palegon 

Thereus palegon (Cramer) 

Lep.: Lycaemidae 129 

thetis, Eucerocoris 

Thrips hawaiiensis Morgan 

Thy.: Thripidae 134 

Thrips tabaci Lindeman Thy.: 

Thripidae 14, 134 

Thrypticus Dip.: 

Dolichopodidae 72 

Thyanta perditor (Fabricius) 

Hem.: Pentatomidae 30


Thysanoplusia orichalcea Uroleucon Hem.: Aphididae walkeri, Psigida 

(~abicius) Lep.: 

Noctuidae 3 1 

tibialis, Thagona 

Tmolus azia (Hewitson) Lep.: 

Lycaenidae 145,162 

Tomaspis entreriana Berg 

Hem.: Cercopidae 143 

Toxoptera graminum, see 

Schizaphis graminum 

trachypterus, Chortogonus 

tredecimpunctatus, 

Rhodobaenus 

triangula, Cicadulina 

triangulifer, Hyalomyodes 

Tricentrus Hem.: 

Membracidae 1 2 1 

Trichogramma Hym.: 

30 

Uroleucon ambrosiae 

(Thomas) Hem.: 

Aphididae 224 

Uroleucon cornpositae 

(Theobald) Hem.: 

Aphididae 188 

Uroleucon solidaginis 


Aphididae 13 

ustulata, Ceresa 

Valtissius Hem.: Lygaeidae 

200 

vanillae, Agraulis 

variegatus, - Zonocerus 

venegasiae, Liriomyza 

Xanthaciura insecta (Loew) 

Dip.: Tephritidae 14,3 1, 

128 

xanthaspis, Exorista 

Xyloplothrips Thy.: 


xylostella, Plutella 

xyonysius Hem.: Lygaeidae 

132 

Xyonysius basalis (Dallas) 

Hem.: Lygaeidae 128 

Xyonysius inaequalis, see 

Xyonysius basalis 

Xyonysius sp. nr ementitus, 

see Xyonysius basalis 

yaseeni, Psyllaephagus 

Trichogrammatidae 1 15 

Trichotaphe, see Dichomeris 

trifolii, Liriomyza 

tripsaci, Dalbu lus 

Tropicomyia theae (Cotes) 


trunculatus, Hypolixus 

turbata, Nymphula 

turbata, Parapoynx 

turrita, Acrida 

undalis, Hellula 

unipunctata, Mythimna 

unipunctata, Pseudaletia 

unipunctatus, Germalus 

upalusalis, Pionea 

urichi, Liothrips 

versicolor, Autoba 

versicolor, Eublemma 

versurella, Coleophora 

Vesiculaphis pieridis Basu 


vibix, Sogatella 

vinitor, Nysius 

virescens, Heliothis 

virescens, Nephotettix 

virgata, Ferrisia 

virginica, Spilosoma 

viridiaeneus, Criptocephalus 

viridigrisea, Austroasca 

viridipennis, Chalcophana 

vitripennis, Hyalopeplus 

vulgaris, Dialeurodes 

zacconius, Chilo 

Zatropis Hym.: Pteromalidae 

130 

zea, Helicoverpa 

zebratus, Acanthoscelides 

Zizeeria knysna (Trimen) 

Lep.: Lycaenidae 22 

Zizeeria krupta (Trimen) 

Lep.: Lycaenidae 22 

Zonocerus variegatus 

(Linnaeus) Ort.: Acrididae 

13,37 

Zulia entreriana, see 

Tomaspis entreriana 

Zygina Hem.: Cicadellidae 

202

General index 

(Bold type indicates the pages alternifolius, Mariscus 

for the main weeds of altissima, Ageratina 

Chapter 4) Amaranthaceae 3,20,21,23, 

abaca (Musa textilis) 1 10 

Acacia mearnsii 1 41, 

143-1 45 

Acacia riparia 132 

Acalitus 125,127,129 

Acalitus adoratus 41,45-48 

Acalypha 100 

Acremonium zonatum 75,76, 

79,82 

adenophora, Ageratum 

adoratus, Acalitus 

Aecidium tithymali 105 

aegyptium, Dactyloctenium 

Aeschynomene sensitiva 161, 

162 

afine, Melastoma 

agerati, Cercospora 

Ageratina altissima 52 

Ageratum adenophora 52 

Ageratum conyzoides 1,3, 

10-16,31,33,50,52,91 

Ageratum riparia 52 

Ageratum vein yellowing 16 

agor 1 10 

Agropyron repens 65 

aguifigay 222 

alang-alang (Imperata 

cylindrica) 37 

alba, Basella 

albida, Mimosa 

albilineans, Xanthornonas 

Albugo bliti 23 

Albugo portulaceae 21 4 

Albugo portulacearum 21 4 

album, Chenopodium 

albus, Amaranthus 

alikbangon 56 

allelopathy 37, 46, 174,230 

Alternanthera philoxeroides 5 

Alternaria 98, 100, 1 16 

Alternaria compacta 23 

Alternaria eichhorniae 77,78 

Alternaria passiforae 189 

Alternaria tenuis 189 

24,216 

amaranthi, Aposphaeria 

amaranthi, Cacodera 

amaranthi, Cercospora 

Amaranthus 20-23 

Amaranthus albus 24 

Amaranthus cruentus 2 1 

Amaranthus dubius 21 

Amaranthus 

hypochondriachus 2 1 

Amaranthus oleosa 22 

Amaranthus spinosus l,3, 

1&24,30,214 

Amaranthus tricolor 22,23 

Amaranthus viridis 2 1-23 

Amblyseius glorius 72 

Amblyseius pederosus 72 

americanum, Pennisetum 

Amphobotrys ricini 100,105 

Anacamseros 2 1 8 

Andropogon annulatus 89 

Andropogon vulgare 94 

Andropogoneae 224 

anemone 2 1 4 

anemone brown ring 21 4 

anemone mosaic 16 

Anhellia niger 41 

anil, Indigofera 

annulatus, Andropogon 

annuus, Helianthus 

anthelminthicum, 

Chenopodium 

Anthemidae 33 

anthocyanin 70 

antidotale, Panicum 

ants 43,45-47,50,52 

anuus, Helianthus 

Aphelenchoides fragariae 15, 

170 

apoda, Phakospora 

apoe apoe 198 

aponapon 198 

Aposphaeria arnaranthi 23 

aquatica, Ipomoea 

ara tanah 104 

Araceae 3, 198 

Arctoteae 33 

arenaria thamesis, 

Meloidogyne 

arenaria, Meloidogyne 

Arundo donax 8 

arvensis, Digera 

arvensis, Mentha 

Ascochyta 225 

Ascochyta portulaceae 2 1 4 

asper, Sonchus 

asperatg, Mimosa 

Aspidosperma ramiflorum 

100 

Aster31,33,214 

aster yellows 32,214 

Asteraceae 3, 12,28-31,33, 

36,38,50,52, 

126,128-130,132 

Astereae 33 

asthma plant (Euphorbia 

hirta) 104 

atramentosa, 

Myriogenospora 

australis, Emex 


inulaefolium 14,30,52 

Avena, 227 

Azolla 7 1 

Azolla caroliniana 205 

Azolla pinnata 1 15,206 

azurea, Eichhornia 

ba bawagan 1 10 

baccata, Wulfla 

baccifera, Melocanna 

bamboo 8 

Bambusa 8,227 

banana 57,63,110,160,180, 

21 4 

banana poka (Pmsifora 

tripartita) 187 

bandotan 12 

banla saet 138 

barley (Hordeum) 8,89,92, 

93


barley stripe mosaic 66 

barnyard grass (Echinochloa 

crus-galli) 62 

Basella alba 22 

bayam duri 20 

bayokibok 62 

bean 88,99 

beda bin 70 

beet curly top 2 14 

Begonia 1 0 1 

benghalensis, Commelina 

bbo chi 198 

Beta vulgaris var. cicla 2 1 1, 

212 

betae f.sp. portulaceae, 

Polymyxa 

bicolor, Cochliobolus 

. bicolor, Helminthosporium 

bicolor, Sorghum 

Bidens 29,3 1 

Bidens mosaic 32 

Bidens mottle 16 

Bidens pilosa l,3,26-33,49, 

52,91,129-131,181 

Bidens witches broom 32 

bidenticola. Uromyces 

bidentis, Cercospora 

biga bigaan 166 

Bignoniaceae 5 1 

Bipolaris indica 24,2 14 

Bipolaris nodulosa, see 


nodulosum 

Bipolaris papendor-i 194 

Bipolaris perotidis 225 

Bipolaris portulaceae, 

see Helminthosporium 

portulaceae 

Bipolaris setariae 86,91 

birds 43,46 

biserrata, Nephrolepis 

bizat 36 

black pepper (Piper nigrum) 

100,122,214 

black wattle (Acacia 

mearnsii) 141,143-1 45 

blady grass (Imperata 

cylindrica) 193 

bliti, Albugo 

blue panic (Panicum 

antidotab) 175 

bb xit 12 

borhg 138 

botrys, Chenopodium 

Brachiaria subquadripara 89 

brachiata, Cercospora 

brAnjiingAn 222 

brassicae, Tylenchorhynchus 

braziliensis, Euphorbia 

Brevipalpus obovatus 15 

Brillantaisia laminum 188 

broad sword fern 

(Nephrolepis biserrata) 

170 

buah tikus 186 

Buddleia variabilis 101 

buffalo grass 180 

bulak manok 12 

bulbosum, Panicum 

bulbous panic (Panicum 

bulbosum) 175 

bullatum, Tolyposporium 

Cacodera amaranthi 23 

cactus 214 

caerulia, Passiflora 

Caesalpiniaceae 139 

cajan, Cajanus 

Cajanus cajan 16 1, 162,188 

Cajanus indicus, see Cajanus 

cajan 

Calacarus 40,41 

Caladium 78 

Calendula 3 1 

Calenduleae 33 

Calliandra selloi 144, 145 

camara, Lantana 

campestiis, Cuscuta 

cancriformis, Triops 

canescens, Cercospora 

capansa, Euphorbia 

capillare, Panicum 

capsicum 91,2 14 

caracasana, Fleischmannia 

caracasana, Wedelia 

Cardueae 33 

Carlineae 33 

caroliniana, Azolla 

carpesioides, Venegasia 

Carthamnus 33 

Caryota 100 

cashew 37 

cassava (Manihot esculenta) 

28,57,63,99,100,181 

Cassia 100 

cassiicola, Corynespora 

Cassytha filiformis 32 

cebbensis, Kordyana 

cenchroides, Pennisetum 

Cercospora 41,167,225 

Cercospora agerati 16 

Cercospora amaranthi 24 

Cercospora bidentis 32 

Cercospora brachiata 24 

Cercospora canescens 145 

Cercospora eupatorii 41 

Cercospora eupatorii- 

ohratii 41 

Cercospora eupatoriicola 41 

Cercospora fusimaculans 225 

Cercospora marsileae 1 16 

Cercospora megalopotamica 

32 

Cercospora piaropi 77,82 

Cercospora portulaceae 2 1 4 

Cercospora rodmanii 69,73, 

75,77,82,83 

Cercospora rottboelliae 226 

Cercosporella dominicana 

214 

Cercosporidium helleri 230, 

23 1 

cereal chlorotic mottle 93 

cespitosa, Euphorbia 

chaetochloae, Puccinia 

chak thorn 198 

Chasmopodium 223 

Chenopodiaceae 2 16 

Chenopodium 2 1,22 

Chenopodium album 22,23 

Chenopodium 

anthelminthicum 22 

Chenopodium botrys 22 

cheroma 126 

chhlong 174 

chi yop luang 148 

chili 2 14 

chili vein banding 2 14 

chinensis, Cuscuta 

Chloridoideae 86 

Chloris gayana 9 1 

chok 198 

Chromolaena 37.52

Chromolaena inulaefolium 52 

Chromolaena iresinoides 14 

Chromolaena ivaefolia 14, 

30,48-52 

Chromolaena jujuensis 47, 

49,50 

Chromolaena laevigata 52 

Chromolaena microstemon 

50 

Chromolaena odorata l,3, 

13, 14,22,24,30,31, 

33,3443,128-132 

Chrysanthemum 3 1,33 

chrysopids 47 

chuntul phnom 1 14 

Cichorium 33 

Cichorium intybus 15 

ciliaris, Digitaria 

Cinchona 13 1,132 

Cineraria 3 1,33 

cinereum, Physarum 

Cionothrix praelonga 38,41 

citrus 22,28,63, 100, 104, 

160,180 

clandestinum, Pennisetum 

Claviceps 175 

clover 2 14 

clover big vein 21 4 

clover fern (Marsilea minuta) 

114 

cb c6t heo 12 

cb hoi 36 

cb l6ng viit 62 

cb m2n Mh 86 

cb 6ng 174 

cb sua l6ng 104 

cb trinh nu m6c 138 

cobbler's pegs (Bidens 

pilosa) 28, 1 30 

cochinchinesis, Rottboellia 

Cochliobolus bicolor 226 

Cochliobolus cymbopogonis 

226 

Cochliobolus heterostrophus 

226 

Cochliobolus lunatus 65 

Cochliobolus nodulosus, see 


nodulosum 

cocoa 37,127,174,18 1 

coconut 28,37,127,139, 

160,174,180,186 

coffee 28,30,3 1,37,63,98, 

100,101,160,180 

Coix lacryma-jobi 8 

Colletotrichium 16, 133,224, 

226 


gloeosporioides f.sp. 

clidemiae 188 


gloesporioides 133,149, 

189 

Colletotrichium graminicola, 

see Glornerella 

grminicola 

Colocasia 78 

Colocasia esculenta 79 

colona, Echinochloa 

coloratum, Panicum 

Commelina 56,57 

Commelina benghalensis 3, 

54-58,91,111 

Commelina difisa 58 

Commelina robusta 57 

Commelinaceae 3,56 

common heliotrope 

(Heliotropiurn europaeum) 

131 

common sensitive plant 

(Mimsa pudica) 160 

compacts, Alternaria 

compressa, Rottboellia 

Coniothyrium 226 

conjugatum, Paspalum 

conoclinii, Puccinia 

conyzoides, Ageratum 

coolah grass (Panicum 

coloratum) 175 

coracana, Eleusine 

cordata, Mikania 

cordata, Pontederia 

cordifolia, Mikania 

Coreopsidae 29,32,33 

Coreopsis 29,33 

corn leaf gall 93,227 

corn stunt 93 

Corticium sasakii 58,91, 1 1 1 

Corynespora cassiicola 1 39, 

1 42 

Cosmos 33 

cotton 20,63,99, 104, 160 

couch grass (Cynodon 

dactylon) 4 

7 General index 293 

coumarin 12 

cowpea 20,142 

crassipes, Eichhornia 

creeping panic grass 

(Panicum repens) 174 

creeping sensitive plant 

(Mimosa invisa) 137,138 

crenata, Marsilea 

Criconemella onoensis 1 1 1 

Criconemella xenoplax 2 1 4 

Crotolaria 10 1 

crotolariae, Phaeotrichoconis 

Croton lobatus 15,91 

crowsfoot grass (Eleusine 

indica) 86 

cruentus, Amaranthus 

crus-galli var. frumentacea, 

Echinochloa 

crus-galli, Echinochloa 

crusgalli, Ustilago 

Ctenopharyngodon idella 72, 

75,76 

cubensis, Diabole 

cucullata, Salvinia 

cucumber 24,214 

cucumber mosaic 24,189, 

214 

cucumeris, Thanatephorus 

Curvularia 133,223,226 

Curvularia cymbopogonis, 

see Cochliobolus 

cymbopogonis 

Cuscuta cmpestris 135 

Cuscuta chinensis 135 

cylindrica, Impera ta 


quinqueseptatum 16, 105 

Cymbopogon 8 

cymbopogonis, Cochliobolus 

cymbopogonis, Curvularia 

Cynara 33 

Cynara scolymus 33 

Cynodon227 

Cynodon dactylon 3,4,15, 

58,64,88-91,111 

cyparissias, Euphorbia 

Cyperaceae 3, 1 10 

Cyperus iria 9 1 

Cyperus rotundus 88,224

294 


Dactyloctenium 86,92,94 

Dactyloctenium aegyptium 

90,92,224 

Dactyloctenium radulans 92, 

94 

dactylon, Cynodon 

Dahlia 33,50 

daoen kaget kaget 160 

dasheen 

(Colocasia esculenta) 79 

daua daua 62 

Davalliaceae 170 

dayflower (Commelina 

benghalensis) 56 

delvii, Heterodera 

den gai 20 

Dendrocalamus 8 


lucknowense 2 14 

destructor, Ditylenchus 

devil weed (Chromolaena 

odorata) 36 

Diabole cubensis 149, 150, 

151, 155, 156 

Diaporthe 226 

dichotomiflorum, Panicum 

Dichotomophthora indica 2 14 

Dichotomophthora lutea, 

see Dichotomophthora 

indica 


portulaceae 21 4 

dictyoides, Drechslera 

difisa, Commelina 

dzfsusa, Marsilea 

Digera arvensis 2 1,22 

Digera mosaic 24 

Digitaria 90,227 

Digitaria ciliaris 91 

Digitaria sanguinalis 63,224 

dilatatum, Paspalum 

Diplodia 226 

dipsaci, Ditylenchus 

distichum, Paspalum 

Ditylenchus destructor 91 

Ditylenchus dipsaci 2 1 4 

djaringan ketul 28 

djawan 62 

Doassansia 167 

dominicana, Cercosporella 

donax, Arundo 

Drechslera dictyoides 65 

Drechslera gigantea 91 

Drechslera hawaiiensis 58 

Drechslera indica, see 

Bipolaris indica 

Drechslera maydis, see 

Cochliobolus 

heterostrophus 

Drechslera setariae, see 

Bipolaris setariae 

dubius, Amaranthus 

dumetorum, Schizostachyum 

duri semalu 138 

2c2ng 70 

echeng padi 166 

echinatum, Pithecoctenium 

Echinochloa 63,227 

Echinochloa colona 63-65, 

89,91,93 

Echinochloa crus-galli 3,57, 

60-66 89,91,111,166 

Echinochloa crus-galli var. 

fiumentacea 62 

Echinochloa glabrescens 93 

Echinochloa oryzicola 63 

Echinochloa pic ta 63.65 

Echinochloa utilis 65 

Echinochloa walteri 65 

Echinopsideae 33 

Echium plantagineum 13 1 

edulis var. flavicarpa, 

Passiflora 

edulis, Passiflora 

eggplant 2 15 

Eichhornia azurea 73, 80, 81 

Eichhornia crassipes 1,3,5, 

68-83,203,205,206 

eichhorniae, Alternaria 

eichhorniae, Flechtmannia 

eichhorniae, Uredo 

ekmaniana, Septoria 

ekor kucing 192 

elephant grass (Pennisetum 

purpureum) 194 

Eleusine 86-88,92,94,226, 

227 

Eleusine coracana 85-89,92, 

93 

Eleusine indica 1,3,5, 15, 

58,84-95,111.224 

Eleusine mosaic 89,93 

Eleusine tristachya 86 

eleusines, Phyllachora 

eleusinis, Melanopsichium 

eleusinis, Ustilago 

Elsinoe 100 

Emex australis 130 

Emex spinosa 130 

encephalitis 71 

Entyloma guaraniticum 32 

Eragrostis 92 

erecta, Tagetes 

ergot 18 1 

erineum 48,129 

erosus, Marsilea 

Erythrina lithosperma 30 

esculenta, Colocasia 

esculenta, Manihot 

etjeng padi 70, 166 

Euchlaena 90 

Eugenia 101 

Eupatorieae 33,52 

eupatorii, Cercospora 

eupatorii, Guignardia 

eupatorii-formosani, 

Pseudocercospora 

eupatorii-odoratii, 

Cercospora 

eupatoriicola, Cercospora 

eupatoriicola, Phomopsis 

eupatoriicola, Phyllosticta 

Eupatorium 13,5 1 

Eupatorium hookerianum, 

see Chromolaena jujuensis 

Eupatorium odoratum, 

see Chromolaena odorata 

Euphorbia 97,99,101 

Euphorbia braziliensis 101 

Euphorbia capansa 100 

Euphorbia cespitosa 101 

Euphorbia cyparissias 99 

Euphorbia geniculata, see 


Euphorbia heterophylla 3,91, 

96-101,104 

Euphorbia hirta 3, 15,99, 

102-106 

Euphorbia hirtella 100 

Euphorbia mosaic 100 

Euphorbia ovalifolia 101 

Euphorbia pilulifera, see 

Euphorbia hirta

Euphorbia prunifolia, see 


Euphorbia pseudovirgata 99 

Euphorbia pulcherrima 99, 

101 

Euphorbiaceae 3,98, 100, 

101,104 

euphorbiae, Uromyces 

europaeum, Heliotropium 

exaltata, Rottboellia 

exigua, Meloidogyne 

Exserohilum paspali 182 

feather Pennisetum 

(Pennisetum polystachion) 

192 

Ficus 77 

filariasis 71 

fili2iformis, Cassytha 

Fimbristylis littoralis, see 


Fimbristylis miliacea 3,57, 

58,91,108-111 

finger millet (Eleusine 

coracana) 85-87 

Flechtmannia eichhorniae 72 

Fleischmannia caracasana 

3 1 

Fleischmannia microstemon 

14 

fluitans, Panicum 

foetida, Passiflora 

fontana, Montia 

foxtail millet (Setaria italica) 

8 

fragariae, Aphelenchoides 

French weed 36 

fujikuroi, Gibberella 

Fusarium 145 

Fusarium moniliforme 226 


elaeidis 24, 41 


passiflorae 188,189 

Fusarium roseum 65 

fusimaculans, Cercospora 

Galinsoga parvi'ora 9 1 

garden spurge (Euphorbia 

hirta) 104 

gatas gatas 104 

gayana, Chloris 

gelang 10 

gelang pasir 210 

gelang susu 104 

gbndong hcok 104 

geniculata, Euphorbia 

gewor 56 

giant sensitive plant (Mimosa 

pigra) 148 

Gibberellafujikuroi, see 

Fusarium moniliforme 

gigantea, Drechslera 

Gigantochloa 8 

glabrescens,. Echinochloa 

glauca, Setaria 

glaucum, Pennisetum 

globe artichoke (Cynara 

scolymus) 33 

Gloeocercospora 194 

gloeosporioides f.sp. 

clidemiae, Colletotrichium 

gloeosporioides, 


Glomerella graminicola 226 

glorius, Amblyseius 

glumae, Pseudomonas 

Glycine max 21 4 

glycines, Heterodera 

goatweed (Ageratum 

conyzoides) 12 

goenda 230 

golasiman 2 10 

goosegrass (Eleusine indica) 

86 

gooseweed (Sphenoclea 

zeylanica) 230 

graminicola, Colletotrichium 

graminicola, Glomerella 

graminicola, Meloidogyne 

grandiflora, Portulaca 

grass carp 

(Ctenopharyngodon idella) 

72 

grass-like fimbristylis 

(Fimbristylis miliacea) 1 10 

Grevillea robusta 101 

grisea, Magnaporthe 

groundnut 20,63,91,93, 104 

210,214 

groundnut rosette 24,32,57, 

93,106,214 


guaraniticum, Entyloma 

Guignardia eupatorii 41 

guinea grass (Panicum 

maximum) 175 

hagonoy 36 

hairy spurge (Euphorbia 

hirta) 104 

hairy wandering jew 

(Commelina benghalensis) 

56 

hapla, Meloidogyne 

Haplosporella passifloridia 

189 

hastata, Monochoria 

hawaiiensis, Drechslera 

hay kai mangda 62 

Heleniae 33 

Heliantheae 33 

Helianthus 24,3 1,33,214 

Helianthus annuus 33 

Helicotylenchus indicus 2 14 

Helicotylenchus multicinctus 

15,57,214 

Heliotropium europaeum 13 1 

helleri, Cercosporidium 

Helminthosporium 65,91,98, 

100 

Helminthosporium bicolor, 

see Cochliobolus bicolor 

Helminthosporium holmii 91 

Helminthosporium maydis 91 


nodulosum 92 

Helminthosporium portulacae 

214 

Helminthosporium rostratum 

194 

Helminthosporium turicum 

65 

Hemphyllium 189 

Heterocentron 

subtriplinenium 12 1 

Heterodera delvii 87,94 

Heterodera glycines 2 1 5 

Heterodera marioni 21 5 

heterophylla, Euphorbia 

heterostrophus, Cochliobolus 

hibiscus yellow vein mosaic 

16,106 

hin nu nive tsu bauk 20

296 


Hirschmaniella 1 1 1, 167,230 

Hirschmaniella oryzae 65 

Hirschmaniella spinicaudata 

9 1 

hirta, Euphorbia 

hirtella, Euphorbia 

holmii, Helminthosporium 

hookeriana, Mikania 

hookerianum, Eupatorium 

Hoplolaimus indicus 2 15 

hordeoides, Pennisetum 

Hordeum 8,227 

hulape 180 

Hydrozetes subornata 20 1 

Hyparrhenia rufa 3 1 

Hypericum perforatum 

angustifolium 130 

hypochondriachus, 

Amaranthus 

idella, Ctenopharyngodon 

Ilex 101 

Impatiens 15 

Imperata cylindrica 5,8,24, 

37,91,186,193 

incognita, Meloidogyne 

Indian rhododendron 

(Melastoma 

rnalabathricum) 120 

indica, Bipolaris 

indica, Dichotomophthora 

indica, Drechslera 

indica, Eleusine 

indicum, Panicum 

indicus, Cajan 

indicus, Hoplolaimus 

indicus, Pseudocephalobus 

Indigofera anil143 

intybus, Cichorium 

inulaefolium, 


Inuleae 33 

invadens, Nosema 

invisa inermis, Mimosa 

invisa, Mimosa 

Ipomoea aquatica 78 

iresinoides, Chromolaena 

iria, Cyperus 

Ischaemum rugosum 225 

italica, Setaria 

itch grass (Rottboellia 

cochinchinensis) 222 

ivaefolia, Chromolaena 

jacobaea, Senecio 

jampang canggah 180 

Japanese millet (Echinochloa 

crus-galli var. 

frumentacea) 62 

javanica, Meloidogyne 

jawg 198 

job's tears (Coix lacryma- 

jobi) 9 

jojoba 2 1 1 

jujuensis, Chromolaena 

jute 63 

ka kiad chrach 166 

ka thok rok 186 

kak phr6k kdam 1 10 

kamplauk 70 

kbet choun 210 

keladi bunting 70 

kelayar 166 

kembang gajah 148 

keruong padi 174 

kiambang besar 198 

kikuyu grass (Pennisetum 

clandestinum) 193 

kirinyu 36 

kolokong kabayo 12 

Kordyana celebensis 58 

krokot 2 10 

kumpai jepang 36 

Lablab purpureus 188 

lablab bean (Lablab 

purpureus) 

lacryma-jobi, Coix 

Lactuca 3 1,33 

Lactuca sativa 33 

Lactuceae 33 

laevigata, Chromolaena 

Lagenaria siceraria 188 

lamaan 62 

laminum, Brillantaisia 

lanceolata, Pontederia 

Lantana 129 

Lantana camara 13,13 1,132 

latex 98,99 

latifolia, Zizania 

Laurus 101 

lawn 4,86, 104, 161, 174, 

181 

legumes 56 

Lemna 71 

Lemna polyrhiza 1 15 

Lemna purpusilla 1 15 

lemon grass (Cymbopogon) 8 

Leptosphaeria 226 

Leptosphaeria proteispora 

182 

lesser fimbristylis 

(Fimbristylis miliacea) 1 10 

lettuce (Lactuca sativa) 30 

Leucaena leucocephala 141, 

142,144,145 

leucocephala, Leucaena 

Lewisia 218 

ligularis, Passifora 

lithosperma, Erythrina 

littoralis, Fimbristylis 

lizards 43,45,46 

lobata, Neurolaena 

lobatus, Croton 

lonchocarpa, Passifora 

love-in-a-mist (Passifora 

foetida) 186 

luc binh 70 

lucerne dwarf 66 

lucknowense, 


lunatus, Cochliobolus 

lutea, Dichotomophthora 

luya luyahan 174 

luzonense, Panicum 

Lyonia ovalifolia 13 

mac co 160 

Macrophomina phaseolina 

100 

macrospora, Sclerophthora 

Madieae 33 

Magnaporthe grisea 87, 92, 

226 

Magnolia pumila 10 1 

mai yah raap yak 148 

maiyaraap thao 138 

mai yarap 160 

mai yarap ton 148 

maize (Zea mays) 5,8,20, 

28,63-65,80,89-93, 

104, 110, 111, 160, 174, 

181,210,222,222-226, 

maize dwarf mosaic 66,93, 

227

maize hoja blanca 227 

maize mosaic 227 

maize nematode 

(Pratylenchus zeae) 91 

maize rayado fine 227 

maize streak 93 

maize stripe 227 

maize stripe tenuivirus 227 

maize white leaf 227 

maize yellow mottle 227 

makahiya 160 

makahiyang lalaki 138 

mala malu 160 

malabathricum, Melastoma 

malaria 7 1 

manatee (Trichechus) 72 

mango 20 

manicata, Passifora 

Manihot esculenta 99 

marioni, Heterodera 

Mariscus alternifolius 24 

Marsilea crenata, see 


Marsilea difisa, 

see Marsilea minuta 

Marsilea erosus, see 


Marsilea minuta 3,112116, 

167 

Marsilea perrieriana, see 

Marsilea crenata 

Marsilea senegalensis, see 


Marsileaceae 3,114 

marsileae, Cercospora 

max, Glycine 

maximum, Panicum 

maydis, Drechslera 

maydis, Helminthosporium 

mayo 104 

mays, Zea 

mearnsii, Acacia 

megalopotamica, Cercospora 

Melanopsichium eleusinis 87, 

92,94 

Melastoma 120 

Melastoma afJine, see 


Melastoma malabathricum 1, 

3,118-123 

Melastoma polyanthum 121 

melastoma (Melastoma 

malabathricum) 120 

Melastomataceae 3, 120, 121 

Melocanna baccifera 8 

Meloidogyne 15,32,91, 162, 

215,224 

Meloidogyne arenaria 15,91, 

215 


thamesis 15 

Meloidogyne exigua 100 

Meloidogyne graminicola 65, 

91,111,167,176,231 

Meloidogyne hapla 32,215 

Meloidogyne incognita 15, 

23,57,91,105,189,215 

Meloidogyne javanica 15,57, 

91,100,106,215 

Meloidogyne oryzae 1 1 1 

melon 99 

Mentha arvensis 100 

Mexican fire plant 

(Euphorbia heterophylla) 

98 

micrantha, Mikania 

microsporida 47,50,52,79 

microstemon, Chromolaena 

microstemon, Fleischmannia 

Mikania 128-1 30,132 

Mikania cordata 126 

Mikania cordifolia 128-1 32 

Mikania hookeriana 13 1 

Mikania micrantha l,3,30, 

33,52,124-135 

Mikania scandens 126 

Mikania trinitaria 128, 13 1, 

132 

Mikania vitifolia 128-1 3 1 

mile-a-minute weed (Mikania 

micrantha) 126 

miliacea, Fimbristylis 

miliaceum, Panicum 

millet 5,20,63,94 

millet panic (Panicum 

miliaceum) 175 

millet, Japanese (Echinochloa 

crus-galli var. 

frumentacea) 62 

Mimosa 

Mimosa albida 144 

Mimosa asperata 148 


Mimosa invisa 1,3,136-145, 

161,162 

Mimosa invisa inermis 139, 

141 

Mimosa pigra I, 3,138, 140, 

142-1 44,146-156,161 

Mimosa pigra berlandieri, 

see Mimosa asperata 

Mimosa pudica l,3, 1 38, 

140-142,144,145, 

158-162 

Mimosa pudica hispida 160 

Mimosa pudica unijuga 160 

Mimosa quadrivalis 144 

Mimosa rixosa 141,144 

Mimosa scabrella 143,145 

Mimosa somnians 141 

Mimosa velloziana 141 

Mimosaceae 3,138,139,148, 

160 

mimosae-invisae, Uredo 

mimosae-pigrae, 

Phloeospora 

mimosine, 149 

minuta, Marsilea 

minutus, Pratylenchus 

Mirabilis 13 

mission grass (Pennisetum 

polystachion) 192 

molesta, Salvinia 

mollissima, Passifora 

m6nhnyin 1 10 

moniliforme, Fusarium 

Monochoria hastata 78 

Monochoria vaginalis 3,78, 

93,115,164-167 

monochoria (Monochoria 

vaginalis) 166 

Montia fontana 2 1 8 

Montia pe$oliata 2 1 8 

multicinctus, Helicotylenchus 

Musa 57 

Mutisieae 33 

mya by it 2 1 0 

Mycovellosiella perfoliata 16, 

4 1 

mye byet 21 0 

myet cho 56 

myet ihi 62 

myet kha 174 

myet ya 222

298 Biological Control of Weeds: Southeast Asian Pro 

myet ya nge 222 

Myriogenospora atramentosa 

182 

Myrothecium roridum 78 

nam nom raatchasee 104 

napier grass (Pennisetum 

purpureum) 194 

needle burr (Amaranthus 

spinosus) 20 

Nephrolepidaceae 170 

Nephrolepis 17 1 

Nephrolepis biserrata 3, 15, 

168-171 

Neptunia 155 

Neptunia oleracea 147, 152, 

155 

Nerium 100 

Neurolaena lobata 13 1 

niger, Anhellia 

nigrum, Solanum 

nodulosa, Bipolaris 

nodulosum, 


nodulosus, Cochliobolus 

Nosema 50,206 

Nosema nr invadens 8 1 

Nosema pilicornis 21 8 

notatum, Paspalum 

novemnerve, Panicum 

novocaledonicus, 

Tetranychus 

oat pseudorosette 66 

oats (Avena sativae) 89,92, 

93 

obovatus, Brevipalpus, 

odorata, Chromolaena 

odoratum, Eupatorium 

oficinarum, Saccharum 

Oidium 149, 162 

oil palm 24,28,37, 127, 160, 

170,180,193 

oleosa, Amaranthus 

oleracea, Neptunia 

oleracea, Portulaca 

oleracea, Spinacia 

onoensis, Criconemella 

ophiuri, Spacelotheca 

ophiuri, Sporisorium 

orai 20 

Orthogalumna terebrantis 69, 

73,75-77,79,80-82 

Oryza 227 

Oryza sativa 8 

oryzae var. commelinae, 

Pyricularia 

oryzae, Hirschmaniella 

oryzae, Meloidogyne 

oryzae, Pyricularia 

oryzicola, Echinochloa 

oryzicola, Eleusine 

ovalifolia, Euphorbia 

ovalifolia, Lyonia 

ovalifolia, Pieris 

overeemi, Sporisorium 

oxysporum f.sp. elaeidis, 

Fusarium 

oxysporum f.sp. passiflorae, 

Fusarium 

paang itik 1 14 

painted spurge (Euphorbia 

heterophylla) 98 

paitan 180 

pak prab 56 

pak vaen 1 14 

paklab 160 

pakpawd 230 

paku larat 170 

palmivora, Phytophthora 

paludosum, Panicum 

panici, 

Parasteneotarsonemus 

paniculatum, Talinum 

Panicum 91,175,177,227 

Panicum antidotale 175 

Panicum bulbosum 175 

Panicum capillare 175 

Panicum coloratum 175 

Panicum dichotomiflorum 65 

Panicum fluitans 177 

Panicum indicum 177 

Panicum luzonense 175 

Panicum maximum 3 1, 175 

Panicum miliaceum 64,90, 

175 

Panicum novemnerve 175 

Panicum paludosum 175 

Panicum repens 3,172-177, 

181 

panicum mosaic 66 

papaya 28,142,160,180 

papendofli, Bipolaris 

Parasteneotarsonemus panici 

175,176 

Paratylenchus 65 

parviflora, Galinsoga 

paspali, Exserohilum 

paspali, Sorosporium 

Paspalum 90, 18 1,227 

Paspalum conjugatum 3,89, 

178-183 

Paspalum dilatatum 18 1 

Paspalum distichum 64,93, 

181 

Paspalum notatum 3 1,89 

Paspalum plicatulum 18 1 

Pasualum scrobiculatum 18 1 

~as>alum vaginatum 1 8 1, 

225 

paspalum (Paspalum 

dilatatum) 18 1 

Passiflora 101, 186-189 

Passiflora edulis 187 

Passiflora edulis var. 

flavicarpa 187, 188 

Passiflora foetida 3,184-189 

Passiflora ligularis 187, 188 

Passiflora lonchocarpa 187 

Passiflora manicata 187 

Passiflora mollissima, see 

Passijlora tripartita 

Passiflora suberosa 187,188 

Passiflora tripartita 185, 187, 

188 

Passifloraceae 3, 186-1 88 

passijlorae, Alternaria 

passifloridia, Haplosporella 

passion vine butterfly 

(Agraulis vanillae) 1 87 

passionfruit chlorotic spot 

189 

passionfruit mosaic 189 

passionfruit ringspot 

potyvirus 189 

passionfruit woodiness 

potyvirus 189 

pasture 4, 12,28,56,86, 139 

181,186 

Paterson's curse (Echium 

plantagineum) 13 1 

pawpaw, see papaya

peanut, see ground nut 

pearl millet (Pennisetum 

glaucum) 94,193,226 

pederosus, Amblyseius 

Pellicularia rolfsii 92 

penniseti, Spacelotheca 

Pennisetum 24, 193, 194, 

2 14,227 

Pennisetum americanum, 

see Pennisetum glaucum 

Pennisetum cenchroides 194 

Pennisetum clandestinum 193 

Pennisetum glaucum 94, 193 

Pennisetum hordeoides 1 92 

Penniseam polystachion 3, 

190-194 


atrichum 192 


polystachion 192 


setosum 192 

Pennisetum polystachyon, see 


Pennisetum purpureum 193 

Pennisetum setosum, see 


Pennisetum typhoideum, see 

Pennisetum glaucum 

pepperwort (Marsilea 

minuta) 1 14 

peregrina, Veronica 

perfoliata, Montia 

perfoliata, Mycovellosiella 

perforatum angustifolium, 

Hypericum 

perotidis, Bipolaris 

perrieriana, Marsilea 

Pestalopsis 149 

Pestalotia 133 

petunia 16 

Phaeoseptoria 226 

phak bia yai 2 10 

phak kbiat 166 

phak khom nam 20 

phak pot 230 

phak, top chawaa 70 

phak waen 1 14 

Phakospora apoda 194 

phaseolina, Macrophomina 

Phaseolus vulgaris 15 

philoxeroides, Alternanthera 

Phloeospora mimosae-pigrae 

147,150,151,155 

Phoenix 100 

Phoma 41,214 

Phoma tropica 24 

Phomopsis 1 49 

Phomopsis eupatoriicola 41 

Phragmites 8 

phti banla 20 

Phyllachora eleusines 92 

Phyllachora sacchari 226 

Phyllostachys 8 

Phyllosticta eupatoriicola 41 

Phyllosticta stratiotes 157 

Physarum cinereum 182 

Phytomonas 105 

Phytophthora palmivora 100, 

105,122,214 

piaropi, Cercospora 

pickerel weed (Pontederia 

cordata) 80 

picta, Echinochloa 

picta, Eleusine 

Pieris ovalifolia 13 

pigeon pea (Cajanus cajan) 

188 

pigra, Mimosa 

pigweed (Portulaca oleracea) 

210 

pilicornis. Nosema 

pilosa, Bidens 

pilosa, Portulaca 

pilulifera, Euphorbia 

pineapple 20, 104, 139, 160, 

170,180 

pineapple yellow spot 14, 16 

pinnata, Azolla 

pis koetjing 138 

pisau pisau 28 

Pistia stratiotes 1,3,5,7 1, 

1 15. 167,196-207 

Pistia virus 1 16, 167 

Pithecoctenium 5 1 

Pithecoctenium echinatum 5 1 

plantagineum, Echium 

plantain 1 11 

plantarii, Pseudomonas 

plicatuhm, Paspalum 

Poaceae 3,62,86,174, 180, 

192,222,225 


poinsettia (Euphorbia 

pulcherrima) 

pokok tjerman 36 

polyanthes, Vernonia 

polyanthum, Melastoma 

Polygonum 13 

polyhedrosis 5 1,187 

Polymyxa betae f.sp. 

portulaceae 21 4 

polyrhiza, Lemna 

polystachion atrichum, 

Pennisetum 

polystachion setosum, 

Pennisetum 

polystachion, Pennisetum 

Pontederia cordata 73,79,80 

Pontederia lanceolata 8 1 

Pontederia rotundifolia 80 

Pontederiaceae 3,70,79,166 

Portulaca 24, 209,218 

Portulaca grandijlora 21 4, 

216,218 

Portulaca oleracea 1,3, 1 5, 

91,208-218 

Portulaca pilosa 21 6 

Portulaca quadrifida 2 16 

portulaceae, Albugo 

portulaceae, Ascochyta 

portulaceae, Bipolaris 

portulaceae, Cercospora 

portulaceae, 


portulaceae, 


Portulacaecae 3,210,216 

portulacearum, Albugo 

potato 15,63, 1 1 1, 167 

potato virus Y 16 

praelonga, Cionothrix 

pratensis, Pratylenchus 

Pratylenchus 57,215 

Pratylenchus minutus ,32, 

215 

Pratylenchus pratensis 1 5,9 1 

Pratylenchus zeae 23,9 1 

proso (Panicum novemnerve) 

175 

proteispora, Leptosphaeria 

prunifolia, Euphorbia 

Pseudocephalobus indicus 23

300 


Pseudocercospora 

eupatorii-formosani 39,41 

Pseudomonas glumae 92 

Pseudomonas plantarii 92 

Pseudomonas solanacearum 

16 

pseudovirgata, Euphorbia 

Puccinia 24, 100, 1 62 

Puccinia conoclinii 16 

Puccinia chaetochloae 194 

Puccinia rottboelliae 

223,226 

Puccinia substriata 194 

pudica hispida, Mimosa 

pudica unijuga, Mimosa 

pudica, Mimosa 

pulcherrima, Euphorbia 

pumila, Magnolia 

pungens, Zoysia 

purpurea, Tephrosia 

purpureum, Pennisetum 

purpureus, Lablab 

purpusilla, Lemna 

purslane (Portulaca oleracea) 

210 

putri malu 1 48 

Pyrenochaeta 227 

Pyricularia 58, 176 

Pyricularia grisea, see 

Magnaporthe grisea 

Pyricularia oryzae 65,92, 

176 

Pyricularia owe var. 

commelinae 58 

quadr$da, Portulaca 

quadripara, Brachiaria 

quadrivalis, Mimosa 

quinqueseptatum, 


Radopholus similis 57, 106, 

215 

radulans, Dactyloctenium 

ragi (Eleusine coracana) 86 

ragwort (Senecio jacobaea) 

131 

ramiflorum, Aspidosperma 

Ranunculus sceleratus 16 

rats 

rau mhc 118 thon 166 

mu Sam 2 1 0 

red gram 22 

r&mb&t& 138 

reniformis, Rotylenchulus 

repens, Agropyron 

repens, Panicum 

Rhizoctonia 65, 133 

Rhizoctonia solani 78, 100, 

167 

rice (Oryza sativa) 5,8, 12, 

20,23,28,56-58,61, 

63-65,71,77,80 88-93, 

98 101,104,110, 

11 1,114,115,149,160 

166,167,174,180 , 186, 

193, 199,202,210,222, 

225 ,226,230 

rice dwarf 66 

rice leaf gall 93,227 

rice orange leaf 93 

rice ragged stunt 93, 167 

rice root rot nematode 91 

rice tungro 66 

rice tungro bacilliform 93 

rice tungro spherical 93 

rice yellow mottle 93 

ricini, Amphobotrys 

riparia, Acacia 

riparia, Ageratum 

rixosa, Mimosa 

robusta, Commelina 

robusta. Grevillea 

rodmanii, Cercospora 

rolfsii, Pellicularia 

rolfsii, Sclerotium 

roridum, Myrothecium 

roseum, Fusarium 

rostratum, Helminthosporium 

Rottboellia 223,227 

Rottboellia cochinchinensis 3, 

92,220-227 

Rottboellia compressa 224, 

225 

Rottboellia exaltata, see 

Rottboellia 

cochinchinensis 

rottboelliae, Cercospora 

rottboelliae, Puccinia 

rottboelliae, Spacelotheca 

rotundifolia, Pontederia 

rotundifolia, Salvinia 

rotundus, Cyperus 

Rotylenchulus reniformis 15, 

23,32,91, 100, 106, 11 1, 

182,215 

rubber 28,37,120,127,139 

160, 170, 174, 180, 193, 

194 

Rudbeckia 3 1 

rufa, Hyparrhenia 

rugosum, Ischaemum 

rumput belulhng 86 

rumput berus 192 

rumput bukit 1 10 

rumput canggah 180 

rumput gajah 192 

rumput go1 kar 36 

rumput jae jae 174 

rumput jurig 192 

rumput kerbau 1 80 

rumput kuning 192 

rumput pait 180 

rumput sambou 86 

rumput tahi ayam 12 

rumput tahi berbau 1 10 

saab sua 36 

sabung sabungan 86 

sacchari, Phyllachora 

Saccharum 224,227 

Saccharum oficinarum 8 

Saccharum spontaneum 227 

saltwater couch (Paspalum 

distichurn or P. vaginatum) 

181 

Salvinia 77, 1 15, 197,202 

Salvinia cucullata 167, 

202, 

Salvinia molesta 5,71,77, 

167,199,202,206 

Salvinia rotundifolia 205 

sambau 62 

sampeas 160 

sanguinalis, Digitaria 

sasakii, Corticium 

sativa, Lactuca 

sativa, Oryza 

sawah 70 

scabrella, Mimosa 

scandens, Mikania 

sceleratus, Ranunculus

Schizostachyum dumetorum 8 

scitaminea, Ustilago 

Sclerophthora macrospora 92 

Sclerotinia sclerotiorum 100 

sclerotiorum, Sclerotinia 

Sclerotium rolfsii 16,92 

scolymus, Cynara 

scrobic (Paspalum 

scrobiculatum) 18 1 

scrobiculatum, Paspalum 

Secale 227 

segan 210 

selloi, Calliandra 

semalu gajah 148 

semhggi 1 14 

sembung rambat 126 

senduduk 120 

Senecio jacobaea 13 1 

Senecioneae 33 

senegalensis, Marsilea 

sensitiva, Aeschynomene 

Septoria 41,47 

Septoria ekrnaniana 41 

sesame 14 

Setaria 227 

Setaria glauca 63 

Setaria italica 8 

setariae, Bipolaris 

setariae, Drechslera 

setosum, Pennisetum 

Siam weed (Chromolaena 

odorata) 36,46 

siceraria, Lagenaria 

silisilihan 230 

silverbeet (Beta vulgaris var. 

cicla) 21 2 

similis, Raabpholus 

sin ngo let kya 86 

sin ngo myet 86 

Skranquia 145 

smao 110 

smao bek kbol62 

smao choeung tukke 86 

Solanaceae 2 16 

solanacearum, Pseuabmonas 

solani, Rhizoctonia 

Solanum nigrum 14,24 

somnians, Mimosa 

Sonchus 130 

Sonchus asper 14 

Sonchus yellow net 32 

Sorghum 5,8,28,63,89,90, 

91,93,104,110,160,210, 

222,224,226,227 

Sorghum bicolor 8 

Sorghum verticillijlorum 93 

Sorosporium paspali 1 82 

sourgrass (Paspalum 

conjugatum) 180 

soybean (Glycine mar) 30,99 

160,222 

soybean mosaic 32 

Spaceloma 100 

Spacelotheca ophiuri, see 

Sporisorium ophiuri 

Spacelotheca penniseti 194 

Spacelotheca rottboelliae 227 

Spanish needle (Bidens 

pilosa) 28 

Sphenoclea zeylanica 3, 167, 

228-231 

Sphenocleaceae 3,230 

spiders 45,46,52 

Spinacia oleracea 22 

spinicaudata, Hirschmaniella 

spinosa, Emex 

spinosus, Amaranthus 

spiny amaranth (Amaranthus 

spinosus) 20 

spiny pigweed (Amaranthus 

spinosus) 20 

Spirodela 204 

spontaneum, Saccharum 

Sporisorium ophiuri 22 1, 

223,227 

Sporisorium overeemi 176 

St John's wort (Hypericum 

perforatum angustifolium) 

130 

stinking passionflower 

(Passijlora foetida) 186 

Straits rhododendron 

(Melastoma 

malabathricum) 120 

stratiotes, Phyllosticta 

stratiotes, Pistia 

strawberry 157 

suberosa, Passzjlora 

subornata, Hydrozetes 

subquadripara, Brachiaria 

substriata, Puccinia 

subtriplinenium, 

Heterocentron 


sugar beet 2 14 

sugarcane (Saccharum 

oficinarum) 5,8,20,63, 

65,89,90,93, 104, 110, 

111, 139, 160, 174, 181, 

193,210,222,226 

sugarcane mosaic 66,93,182 

sugarcane streak 93 

sunflower (Helianthus anuus) 

33 

swamp panic (Panicum 

paludosum) 175 

sweet potato 30,57 

Tageteae 33 

Tagetes erecta 24 

Talinum paniculatum 2 1 8 

Talinum triangulare 21 8 

tapah itik 1 14 

tapioca mosaic 16, 106 

taro (Colocasia) 57,63,71, 

79,110 

tarragon 2 1 4 

tawbizat 36 

tea 28, 37, 104, 127, 139, 

160,174,180 

teak 37,127 

tenuis, Alternaria 

Tephrosia purpurea 37 

terebrantis, Orthogalumna 

termites 43 

Tetranychus 140 

Tetranychus novocaledonicus 

23 

Tetranychus urticae 15,100 

Thanatephorus cucumeris 

167 

Tiboochina urvilleana 121 

timun padang 186 

tithymali, Aecidium 

toads 45 

tobacco 28,30,63,91,214 

tobacco broad ring spot 21 4 

tobacco bunchy top 24 

tobacco etch 21 4 

tobacco leaf curl 16, 106 

tobacco mosaic 24,2 14 

tobacco streak 2 14 

Tolyposporium bullatum 66 

tomato91, 100, 111, 142 

tomato leaf curl 1 6, 106


tomato spotted wilt 32 

tontrem khet 36 

torpedo grass (Panicum 

repens) 174 

Tradescantia 57 

Tragia volubilis 101 

translucens, Xanthomonas 

triangulare, Talinum 

Trichechus 72 

tricolor, Amaranthus 

tricophora, Ustilago 

trinh nu nhon 148 

trinitaria, Mikania 

Triops cancriformis 1 67 

tripartita, Passiflora 

tristachya, Eleusine 

Triticum 8,227 

tropica, Phoma 

tuk das khla thom 104 

tungro 93 

turicum, Helminthosporium 

Turneraceae 187 

Tylenchorhynchus brassicae 

215 

typhoideum, Pennisetum 

ulam tikus 126 

ulasiman 21 0 

urd bean yellow mosaic 16, 

106 

Ureab eichhorniae 82 

Ureab mimosae-invisae 145 

Uromyces bidenticola 32 

Uromyces euphorbiae 100 

urticae, Tetranychus 

urvilleana, Tiboochina 

Ustilago 92 

Ustilago crusgalli 66 

Ustilago eleusinis, see 

Melanopsichium eleusinis 

92 

Ustilago sciraminea 227 

Ustilago tricophora 66 

utilis, Echinochloa 

utilis, Eleusine 

vaginalis, Monochoria 

vaginatum, Paspalum 

Vanda 15 

variabilis, Buddleia 

vegetables 8,20,28,56,63, 

104,181,193,210 

velloziana, Mimosa 

Venegasia carpesioides 3 1 

Verbena 16 

Verbesina 3 1 

Vemonieae 33 

Veronica peregrina 167 

verticilliflorwn, Sorghum 

viridis, Amaranthus 

vitifolia, Mikania 

volubilis, Tragia 

vulgare, Andropogon 

vulgaris var. cicla, Beta 

vulgaris, Phaseolus 

walteri, Echinochloa 

water clover (Marsilea 

minuta) 1 1 3-1 15 

water hyacinth (Eichhornia 

crassipes) 70 

water lettuce (Pistia 

stratiotes) 198 

Wedelia caracasana 1 4 

wheat (Triticum) 58,65, 

89-93,111,214 

wheat rosette 93 

wheat streak mosaic 66 

white amur 

(Ctenopharyngdon idella) 

72 

wild passionfruit (Passiflora 

foetida) 186 

witchgrass (Panicum 

capillare) 175 

Wulfia baccata 14 

xB b6ng 230 

Xanthium 15 

Xanthomonas albilineans 182 

Xanthomonas translucens 66 

xenoplar, Criconemella 

ya hep 180 

ya plong 62 

ya sap raeng 12 

ya tabsua 12 

yaa khaehyon chop k92 

yaa prong khaai 222 

yaa yaang 98 

yah chan ah kat 174 

yah chanagard 174 

yah koen jam khao 28 

yah sua mop 36 

yah teenka 86 

ye padauk 70 

Zea 224,227 

Zea mays 8 

zeae, Pratylenchus 

zeylanica, Sphenoclea 

Zinnia 16,3 1,33 

Zinnia yellow net 16 

zonatum, Acremonium 

Zoysia pungens 18 1

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