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Biological Control of Weeds:<br />
Southeast Asian Prospects<br />
D.F. Waterhouse<br />
(ACIAR Consultant in Plant Protection)<br />
ACIAR<br />
(Australian Centre for International Agricultural Research)<br />
Canberra<br />
AUSTRALIA<br />
1994
The Australian Centre for Intemational Agricultural Research (ACIAR) was established in June 1982 by an<br />
Act of the Australian Parliament. Its primary mandate is to help identify agricultural problems in developing<br />
countries and to commission collaborative research between Australian and developing country researchers<br />
in fields where Australia has special competence.<br />
Where trade names are used this constitutes neither endorsement of nor discrimination against any product<br />
by the Centre.<br />
ACIAR MONOGRAPH SERIES<br />
This peer-reviewed series contains the results of original research supported by ACIAR, or deemed relevant<br />
to ACIAR's research objectives. The series is distributed internationally, with an emphasis on the Third<br />
World.<br />
O Australian Centre for Intemational Agricultural Research<br />
GPO Box 157 1, Canberra, ACT 260 1.<br />
Waterhouse, D.F. 1994. Biological Control of Weeds: Southeast Asian Prospects<br />
ACIAR Monograph No. 26, vi + 302pp., 27 figs., 28 maps<br />
ISBN . 186320099 1<br />
Typset in 11/13 Times using a Macintosh IIvx running Quark XPress by K & B Publications<br />
Printed by Bmwn Prior Anderson Pty. Ltd.
Foreword<br />
Contents<br />
Abstract<br />
Estimation of biological control prospects<br />
Introduction<br />
Target weeds<br />
Ageratum conyzoides<br />
Amaranthus spinosus<br />
Bidens pilosa<br />
Chromolaena odorata<br />
Commelina benghalensis<br />
Echinochloa crus-galli<br />
Eichhornia crassipes<br />
Eleusine indica<br />
Euphorbia heterophylla<br />
Euphorbia hirta<br />
Fimbristylis miliacea<br />
Marsilea minuta<br />
Melastoma rnalabathricum<br />
Mikania micrantha<br />
Mimosa invisa<br />
Mimosa pigra<br />
Mimosa pudica<br />
Monochoria vaginalis<br />
Nephrolepis biserrata<br />
Panicum repens<br />
Paspalurn conjugatum<br />
PassiJlora foetida<br />
Pennisetum polystachion<br />
Pistia stratiotes<br />
Portulaca oleracea<br />
Rottboellia cochinchinensis<br />
Sphenoclea zeylanica<br />
References<br />
Index of scientific names of insects<br />
General index
Foreword<br />
From its very beginning in 1982 ACIAR has been a strong supporter of biological control<br />
as a sustainable and environmentally friendly alternative to the steadily growing use of<br />
pesticides. This alternative has achieved great success in regions of the world<br />
(e.g. Australia, New Zealand, Oceania, California) where many of the major insect pests<br />
and weeds have been introduced from outside the region. Although a smaller proportion<br />
of the major weeds in Southeast Asia are introduced than in many other regions, a recent<br />
survey commissioned by ACIAR (Waterhouse 1993a) identified 28 major weeds that<br />
merited evaluation as possible targets for biological control. Even if only half of these<br />
weeds proved to be attractive targets, this number would require several decades of<br />
research, major resources in personnel and equipment and strong support within the<br />
region.<br />
The aim of the present volume is to summarise for the major exotic weeds of agri-<br />
culture in Southeast Asia what is known about their natural enemies and the prospects for<br />
classical biological control. The book is intended to serve two purposes. Firstly, to facili-<br />
tate, for the countries of the region, the selection of promising, individual or collabora-<br />
tive, priority weed targets. Secondly, to provide donor agencies with an overall perspec-<br />
tive of the region's major exotic weed problems and prospects for their amelioration; and<br />
thus to aid in the selection of projects for support that are best suited to their terms of ref-<br />
erence.<br />
It is hoped that it may be possible in the near future to produce a companion volume<br />
dealing with major arthropod pests exotic to Southeast Asia.<br />
G.H.L. Rothschild<br />
Director<br />
Australian Centre for International<br />
Agricultural Research, Canberra
Abstract<br />
Biological control programs have already been mounted in some region of the world<br />
against 6 of the 28 major weeds that are exotic to Southeast Asia. Substantial or partial<br />
success has been achieved in one or more countries for all of these except Mikania<br />
micrantha, which is still under investigation. A substantial amount of information on<br />
their natural enemies in the region where the weeds evolved is available on all 6. This is<br />
in stark contrast with the situation for most of the remaining 22 weed species. Indeed, for<br />
more than half of these, so little relevant information is available that it is not possible to<br />
evaluate the chances of mounting a successful program. For this group of weeds the first<br />
step would be a survey in the centre of origin of the weed. It is probable that surveys<br />
could be mounted simultaneously of several candidate weeds in the same region of the<br />
world (e.g. Central America or Tropical Africa). The very minimum period for a prelimi-<br />
nary survey would be several weeks in both spring and late summer. When the organisms<br />
collected had been identified by taxonomists a decision would be facilitated on possible<br />
follow-up surveys.<br />
On the basis of available information there are good to excellent prospects for<br />
reducing, in at least some parts of the region, the weediness of the following:<br />
Chromolaena odorata<br />
Eichhornia crassipes<br />
Mimosa invisa<br />
Mimosa pigra<br />
Pistia stratiotes<br />
Portulaca oleracea<br />
There are also good reasons for believing that there will prove to be valuable natural<br />
enemies for the following:<br />
Ageratum conyzoides<br />
Amaranthus spinosus<br />
Bidens pilosa<br />
Eleusine indica<br />
Melastoma malabathricum<br />
Mikania micrantha<br />
There is insufficient information yet available on the remaining 15 weeds to attempt<br />
to evaluate their prospects for classical biological control.
2 Estimation of biological control prospects<br />
Weed Rating<br />
Ageratum conyzoides<br />
Amaranthus spinosus<br />
Bidens pilosa<br />
Chromolaena odorata<br />
Commelina benghalensis<br />
Cynodon dactylon<br />
Echinochloa crus-galli<br />
Eichhornia crassipes<br />
Eleusine indica<br />
Euphorbia heterophylla<br />
Euphorbia hirta<br />
Fimbristylis miliacea<br />
Marsilea minuta<br />
Melastoma malabathricum<br />
Mikania micrantha<br />
Mimosa invisa<br />
Mimosa pigra<br />
Mimosa pudica<br />
Monochoria vaginalis<br />
Nephrolepis biserrata<br />
Panicum repens<br />
Paspalum conjugatum<br />
PassiJlora foetida<br />
Pennisetum polystachion<br />
Pistia stratiotes<br />
Portulaca oleracea<br />
Rottboellia cochinchinensis<br />
Sphenoclea zeylanica<br />
Any<br />
Family biological<br />
control<br />
successes?<br />
Asteraceae<br />
Amaranthaceae<br />
Asteraceae<br />
Asteraceae<br />
Commelinaceae<br />
Poaceae<br />
Poaceae<br />
Pontederiaceae<br />
Poaceae<br />
Euphorbiaceae<br />
Euphorbiaceae<br />
C yperaceae<br />
Marsileaceae<br />
Melastomataceae<br />
Asteraceae<br />
Mimosaceae<br />
Mimosaceae<br />
Mimosaceae<br />
Pontederiaceae<br />
Nephrolepidaceae<br />
Poaceae<br />
Poaceae<br />
Passifloraceae<br />
Poaceae<br />
Araceae<br />
Portulacaceae<br />
Poaceae<br />
Sphenocleaceae<br />
Attractiveness<br />
as a target in<br />
SE Asia<br />
unsuitable
4 Biological Control of Weeds: Southeast Asian Prospects<br />
3 Introduction<br />
Waterhouse (1 993a) published information, collated from agricultural and weed experts<br />
in the 10 countries of Southeast Asia, on the distribution and importance of their major<br />
weeds in agriculture. Ratings were supplied on the basis of a very simple system:<br />
+++ very widespread and very important<br />
++ not widespread but of great importance where it occurs<br />
+ important only locally<br />
present, but not an important pest<br />
The advantages and limitations of this system are discussed by Waterhouse (1993a). Of<br />
232 weeds nominated, 140 were rated as highly important, and a subset of 40 particularly<br />
SO.<br />
The focus of the present work is on the possibilities for classical biological control<br />
of those of this subset of 40 that evolved outside Southeast Asia. The assumption is that<br />
many of these have been introduced without some of the organisms that help to control<br />
them where they evolved. The chances are very remote indeed, for weeds that evolved in<br />
Southeast Asia, of introducing sufficiently host-specific organisms from outside the<br />
region. Nevertheless, it is possible that useful organisms present in, say, Thailand or<br />
Myanmar may not be present in all of the islands constituting the Philippines or<br />
Indonesia (or vice versa) and this possibility should be borne in mind.<br />
The origin of 12 of the subset of 40 major weeds is believed to be Southeast Asia, or<br />
close by, and these have been excluded from consideration at this stage. The remaining<br />
28 species, 27 of which are treated here, are either known to have evolved in the<br />
Americas or Africa or are postulated to have evolved in both Asia and Africa. This latter<br />
group is considered because the possibility exists that useful organisms at the African end<br />
of the range may not yet have extended their distribution into all of Southeast Asia.<br />
The 28th species, couch grass, Cynodon dactylon, has not been dealt with because,<br />
in many situations, such as lawns and some pastures, it is regarded as a highly desirable<br />
species. Biological control agents would not distinguish between these situations and the<br />
many others where it is a serious weed, so other control measures must be employed in<br />
the latter instances.<br />
Of course, it is not to be expected that all of any one country's top 20 or even top 10<br />
exotic weeds will necessarily be included in this regional priority list. Indeed, at least<br />
some of those omitted might well merit the production of additional dossiers if they are<br />
of such importance locally that resources for a program would be likely to achieve a very<br />
high priority for a particular country. ACIAR would be interested to hear of weeds that<br />
might be considered in this category.<br />
It is not so long ago that Wilson (1964) pointed out that no insects had yet been used<br />
for the biological control of aquatic weeds and that it was not clear "whether in the aquat-<br />
ic environment there exists a sufficient development of that monophagy in phytophagous<br />
insects that has been the main foundation for the biological control of weeds on land". He
Introduction 5<br />
referred to the opinion of Brues (1946) that aquatic insects show little host specificity, but<br />
warned that this view might be the result of lack of information and recommended an<br />
extension of research in this general field. In the intervening 30 years, research on four<br />
major water weeds of South American origin has yielded success and even spectacular<br />
success with the following: Salvinia molesta, Eichhornia crassipes, Alternanthera<br />
philoxeroides and Pistia stratiotes (Room 1993).<br />
It is very probable that a parallel can be drawn between the situation with water<br />
weeds in 1964 and the "conventional wisdom" of today that grassy weeds are unsuitable<br />
targets for classical biological control because of the danger to many major world crops<br />
that also belong to the family Poaceae e.g. rice, wheat, maize, sorghum, millet, sugar-<br />
cane. However, it would be very strange indeed if host specialisation occurred widely in<br />
insects attacking all other plant families, but not amongst those attacking the very large<br />
number of grasses. In view of the fact that 10 of the 18 world's worst weeds are grasses<br />
(Holm et al. 1977) and eight of the 28 major exotic weeds in Southeast Asia are also<br />
grasses (Waterhouse 1992, 1993a), it is evident that the time is long overdue for a<br />
detailed study of the natural enemies of these grasses in the regions where they evolved.<br />
This theme is mentioned further below, in particular in the discussion on Eleusine indica.<br />
For any biological control organisms to be approved for introduction into Southeast<br />
Asia against weedy grasses they would need to be sufficiently specific that they would<br />
not cause economic damage to the crop grasses listed in table 3.1. This list refers to<br />
Thailand, but is believed to be much the same as that for other Southeast Asian countries.<br />
It does not, however, include pasture species. A number of useful grasses are also har-<br />
vested from the wild and some may have to be considered also, although there are impor-<br />
tant weeds (e.g. Zmperata cylindrica) amongst them. There are, of course, many addition-<br />
al crop grasses of importance outside the region, but of little or no importance in most or<br />
all of Southeast Asia. They would certainly have to be taken into consideration in other<br />
regions of the world.<br />
The successful biological control of a weed presents a special problem, seldom<br />
shared by the control of an insect pest, namely that some other plant, perhaps even a<br />
weed that is more difficult to control by other means, will spread to occupy the space<br />
vacated. Reduction to the greatest possible extent of the density of a weed is desirable in<br />
situations such as pastures or national parks. In many other situations, however, all that<br />
may be required is a significant reduction in seeding (for annuals) or in competitiveness<br />
(for annuals and perennials) so that the weed no longer has an opportunity of becoming<br />
dominant and thus, when necessary, is more readily controlled by cultural or other mea-<br />
sures. Thus, even partial biological control (leading to the weed becoming less aggres-<br />
sive) provides desirable plant species with the opportunity to compete more successfully<br />
for sunlight and nutrients and may be of significant value.<br />
Another problem is that many weeds display a good deal of variability throughout<br />
their distribution, resulting in part from polyploidy, hybridisation with closely related<br />
species and other genetic modifications. The taxa thus produced may not be equally sus-<br />
ceptible to natural enemies, so it is desirable, where possible, to match-tb-wlth ixa<br />
encountered in the surveys in the area of origin of the weed. It may also be necessary to
6 Biological Control of Weeds: Southeast Asian Prospects<br />
seek expert taxonomic advice at an early stage, perhaps involving electrophoretic, DNA<br />
and other studies, particularly when commencing a project on a weed that has not yet<br />
been the target of a biological control investigation.<br />
The summary accounts presented are designed to enable a rapid review to be made<br />
of (i) the main characteristics of the major weeds of agriculture that are believed to be<br />
exotic to part or all of Southeast Asia, (ii) what is known of their natural enemies and<br />
(iii) prospects for reducing their weediness by classical biological control.<br />
The material on weed characteristics draws heavily on the publications by Barnes<br />
and Chan (1990), Holm et al. (1977), Noda et al. (1985) and Soerjani et al. (1987).<br />
Additional information is available from these sources, including detailed botanical<br />
descriptions, vernacular names, biology, agricultural importance and herbicidal control.<br />
I am particularly grateful to the University of Hawaii Press for permission to draw<br />
on 21 of the illustrations in its publication 'The World's Worst Weeds' by Holm et al.<br />
(1977) to Ancom Berhad, Malaysia (Barnes and Chan 1990) and the Director of<br />
BIOTROP Indonesia (Soerjani et al. 1987) to draw on 2 and 3 illustrations respectively<br />
from their publications and to the Division of Entomology CSIRO for permission to use<br />
illustration 4.16. The figures have been slightly amended by the omission of inserts that<br />
are mainly of taxonomic interest. Acknowledgement appears on each of the illustrations<br />
used.<br />
In most instances four databases were searched for relevant information:<br />
AGRICOLA (Bibliography of Agriculture) 1 970+<br />
BIOSIS (Biological Abstracts) 1989+<br />
CAB (Commonwealth Agricultural Bureaux) 1984+<br />
DIALOG (Biological Abstracts) 1959+<br />
1 many cases abstracting journals and other sources published prior to the above<br />
cominencement dates were also searched. Useful information was also obtained by<br />
serendipity from these and other references and from unpublished records. Nevertheless,<br />
in many cases the search cannot be described as exhaustive. Even more relevant, howev-<br />
er, than attempting an exhaustive search would be a fresh, detailed field survey targeted<br />
on the known (or presumed) area of origin of the weed. In any event, in most instances a<br />
preliminary investigation would be highly desirable in the area of origin of a weed before<br />
deciding whether or not to embark upon a major project. Several such surveys might well<br />
be carried out simultaneously where more than one weed occurs in the same general<br />
region. Indeed, it is strongly recommended that a pre-project activity be funded to carry<br />
out such surveys, with special reference to selected weeds of major importance in<br />
Southeast Asia.<br />
Surveys of this nature are particularly important, since the amount of useful, pub-<br />
lished information on arthropods or other organisms attacking the target weeds is, in gen-<br />
eral, inadequate to serve as a basis for a sound decision. Although acceptable host speci-<br />
ficity is required for classical biological control, it is possible that some of the less specif-<br />
ic fungi listed might be developed for use as bioherbicides.<br />
In addition to surveys in the region of origin of the weed(s) it will also be necessary<br />
to survey the weed(s) in the country or countries where biological control is to be
Introduction 7<br />
attempted. This is to indicate whether any of the organisms that might be considered for<br />
introduction are already present.<br />
The species treated are drawn from tables 10 and 11 of 'The Major Arthropod Pests<br />
and Weeds of Agriculture in Southeast Asia: Distribution, Importance and Origin'<br />
(Waterhouse 1993a). It is quite possible that additional weeds rating highly in these<br />
tables will prove to be exotic to Southeast Asia (or significant parts of it) and, alternative-<br />
ly, that some considered to be exotic will, on further evidence, be shown to have evolved<br />
in the region.<br />
The natural enemies most commonly involved in classical biological control of<br />
weeds have been arthropods, although there is a growing interest in, and a few striking<br />
successes with, fungi. Because there is a considerable lack of uniformity in the names of<br />
many of the insects involved, a separate index is included listing the preferred scientific<br />
names. These have been used in the text, replacing those used by the authors quoted. On<br />
the other hand, with few exceptions the names used for fungi, bacteria, nematodes and<br />
viruses are those of the authors quoted, although it is probable that some names have<br />
been changed since they were used. Where the name of a weed or an insect given in a<br />
publication is no longer preferred by taxonomists, the superseded name, x, is shown thus<br />
(= x), but this usage is not intended to convey any other taxonomic message. Indeed, the<br />
superseded name may still be valid, but simply not applicable to the particular species<br />
referred to by the author.<br />
I am most grateful for assistance from many colleagues during the preparation of<br />
this book. It is not possible to name them all, but special thanks are due to Dr B.<br />
Napompeth (Thailand), Dr R. Muniappan (Guam), C.J. Davis (Hawaii) and, in Australia,<br />
Dr I.W. Forno, Dr K.L.S. Harley, M.H. Julien, Dr K.R. Norris, J. Prance, Dr D.P.A.<br />
Sands, Dr A.J. Wapshere and A.D. Wright of CSIRO and Dr R.E. McFadyen<br />
(Queensland Department of Lands). Many others who have contributed unpublished<br />
information are acknowledged at appropriate places in the text.<br />
Valuable advice on taxonomic problems has been received from a number of col-<br />
leagues in the Division of Entomology, CSIRO, Canberra, including Dr M. Carver<br />
(Hemiptera), Dr P. Cranston (Diptera), E.D. Edwards (Lepidoptera), Dr I.D. Naumann,<br />
Dr K.H.L. Key (Orthoptera), T. Weir (Coleoptera) and Dr E.C. Zimmerman<br />
(Curculionidae).<br />
Continuing warm support has been provided by Dr P. Ferrar, Research Program<br />
Coordinator, Crop Scienqs, ACIAR, Canberra.<br />
It is a pleasure to acknowledge the expert assistance of Mrs A. Johnstone<br />
(Ms A. Ankers) in converting my manuscripts into presentable form; and also of Mrs S.<br />
Smith and C. Hunt for assistance with the illustrations.<br />
It would not have been possible to continue with these biological control activities<br />
in deep retirement without the support, forbearance and encouragement of my wife, to<br />
whom particular thanks are due.
8<br />
Biological Control of Weeds: Southeast Asian Prospects<br />
Table 3.1 Grasses (other than pasture species) that are important in Thailand.<br />
A. Crop Grasses Importance<br />
Bambusa spp.<br />
Coix lacryma-jobi<br />
Cymbopogon spp.<br />
Hordeum spp.<br />
Oryza sativa<br />
Saccharum oficinarum<br />
Setaria italica<br />
Sorghum bicolor<br />
Triticum spp.<br />
Zea mays<br />
Zizania latifolia<br />
B. Grasses harvested from the wild<br />
Arundo donax<br />
Dendrocalamus spp.<br />
Gigantochloa spp.<br />
Imperata cylindrica<br />
Melocanna baccifera<br />
Phragmites spp.<br />
Phyllostachys spp.<br />
Schizostachyum dumetorum<br />
bamboo, construction, furniture,<br />
paper<br />
job's tears, cereal<br />
lemongrasses, flavourings<br />
barleys<br />
rice<br />
sugar cane<br />
foxtail millet<br />
sorghum<br />
wheats<br />
maize<br />
vegetable<br />
giant reed, cane<br />
weaving, vegetables<br />
construction, furniture<br />
paper, roof thatch<br />
paper, furniture, food<br />
reeds, thatch, mats<br />
furniture, vegetable<br />
rope
Target weeds<br />
Ageratum conyzoides<br />
Amaranthus spinosus<br />
Bidens pilosa<br />
Chromolaena odorata<br />
Commelina benghalensis<br />
Echinochloa crus-galli<br />
Eichhornia crassipes<br />
Eleusine indica<br />
Euphorbia heterophylla<br />
Euphorbia hirta<br />
Fimbristylis miliacea<br />
Marsilea minuta<br />
Melastoma malabathricum<br />
Mikania micrantha<br />
Mimosa invisa<br />
Mimosa pigra<br />
Mimosa pudica<br />
Monochoria vaginalis<br />
Nephrolepis biserrata<br />
Panicurn repens<br />
Paspalum conjugatum<br />
Passiflora foetida<br />
Penniseturn polystachion<br />
Pistia stratiotes<br />
Portulaca oleracea<br />
Rottboellia cochinchinensis<br />
Sphenoclea zeylanica
10 Biological Control of Weeds: Southeast Asian Prospects<br />
Ageratum conyzoides<br />
(after Holm et a/. 1977)
Map 4.1 Ageratum conyzoides<br />
4.1 Ageratum conyzoides 11<br />
Ageratum conyzoides<br />
As a member of the Asteraceae, it would be expected that Ageratum conyzoides would<br />
have many natural enemies attacking it in its area of origin in Tropical America.<br />
However, no study has been made and virtually nothing is known of the situation there.<br />
Elsewhere it is attacked by a range of insects, nematodes, fungi and viruses, but almost<br />
all have a very wide host range and are not suitable as biological control agents.<br />
Surveys in Tropical America would be necessary to provide data on which<br />
prospects for its biological control could be evaluated.
12 Biological Control of Weeds: Southeast Asian Prospects<br />
4.1 Ageratum conyzoides L.<br />
Asteraceae<br />
goatweed, ageratum; bandotan (Indonesia), rumput tahi ayam (Malaysia), bulak<br />
manok, kolokong kabayo (Philippines), ya tabsua, ya sap raeng (Thailand),<br />
co c~lt heo, bo xit (Vietnam)<br />
Rating<br />
+++ Myan, Thai<br />
17 ++ Msia, Sing, Phil<br />
+ Laos, Camb, Viet, Brun, Indo<br />
Origin<br />
Tropical America.<br />
Distribution<br />
Pantropical; also in the subtropics and extending into temperate areas from latitude 30°N<br />
to 30's. Widespread in SE Asia. Present in Java prior to 1860.<br />
Characteristics<br />
Ageratum conyzoides is a self pollinated, C3, annual herb. It is erect, often branched,<br />
sometimes decumbent and ranges up to 1.2 m at flowering. Its flowers are light blue,<br />
white or violet and its leaves and stems are hairy.<br />
Importance<br />
A. conyzoides occurs in both light and heavy soils in moister areas in agricultural land,<br />
waste land, roadsides, plantations, pastures and upland rice fields. It may produce<br />
40 000 or more seeds per plant and these are mainly spread by wind and water. They<br />
germinate readily and the life cycle can be completed in less than 2 months. A. cony-<br />
zoides is one of about 300 species in the genus, all of which originated in the Americas.<br />
Goatweed is important in 46 countries in 36 crops and is troublesome in plantations<br />
after grasses have been suppressed (Holm et al. 1977). It is a rapidly colonising, vigor-<br />
ously growing weed in a wide variety of arable crops in which thick carpets of A. cony-<br />
zoides compete strongly for nutrients and moisture. When a stand is destroyed another<br />
rapidly takes its place. It is suspected of poisoning cattle, but this is not confirmed from<br />
Australia. It was rated 19th of the World's Worst Weeds by Holm et al. (1977), as equal<br />
15th in Southeast Asia (Waterhouse 1993a) and 15th in the Oceanic Pacific<br />
(waterhouse unpub.).<br />
Its crushed leaves smell strongly of coumarin and are used as a styptic for wounds,<br />
also for sores, skin diseases, eye inflammation and lung problems (Gonzalez et al. 1991).<br />
It is sometimes used as cut flowers in the home.
Natural enemies<br />
4.1 Ageratum conyzoides 13<br />
Although A. conyzoides is listed by Holm et al. (1977) as a weed in some crops in<br />
Central and South America, it is significant that nowhere in that region (unlike the rest of<br />
the tropical world) is it regarded as a serious or a principal weed. From this it might be<br />
inferred that natural enemies might be controlling its abundance. However, so little infor-<br />
mation on natural enemies (Tables 4.1.1 to 4.1.3) was obtained from the databases<br />
searched that it is not possible to substantiate this claim. Almost all of the records are<br />
from outside its area of origin and one (the agromyzid fly, Melanagromyza metallica) is<br />
known to have a narrow host range. However, M. metallica is already widespread. In<br />
addition to India, it is known also from many places including Taiwan, Philippines,<br />
Vietnam, Thailand, Indonesia, Melanesia, Papua New Guinea, Solomon Is, Micronesia,<br />
Australia and Africa. It lays its eggs on the apical part of the stem. The larva bores into<br />
the pith region, gradually extending towards the root and the final instar larva cuts an exit<br />
hole at the base of the stem. Mines may extend into the roots and pupae are often present<br />
at about ground level in the mines (Singh and Beri 1973).<br />
If A. conyzoides is considered to be an important target it will be necessary to sur-<br />
vey for organisms attacking it in Central America and northern South America.<br />
Table 4.1.1 Natural enemies of Aaeratum convzoides: insects and mites.<br />
Species Location Other hosts References<br />
INSECTS<br />
Orthoptera<br />
ACRlDlDAE<br />
Zonocerus Nigeria<br />
variegatus<br />
Hemiptera<br />
APHlDlDAE<br />
Aphis craccivora<br />
Aphis gossypii<br />
Aphis spiraecola Java<br />
(= A. nigricauda)<br />
Aulacorthum<br />
solani<br />
Brachycaudus<br />
helichrysi<br />
Capitophorus<br />
hippophaes<br />
Hyperomyzus<br />
carduellinus<br />
Myzus ormtus<br />
Myzus persicae<br />
Neomasonaphis<br />
(- Masonaphis)<br />
amphalidis<br />
Uroleucon<br />
(= Macrosiphum)<br />
solidaginis<br />
Vesiculaphis pieridis India<br />
Chromolaena odorata,<br />
Lantana camara<br />
many<br />
many<br />
many<br />
many<br />
many<br />
Eupatorium, Mirabilis,<br />
Polygonum<br />
many<br />
many<br />
many<br />
Lyonia ovalifolia,<br />
Pieris ovalifolia<br />
Raychaudhuri 1983<br />
Raychaudhuri 1983<br />
Patch 1939,<br />
Raychaudhuri 1983<br />
Raychaudhuri 1983<br />
Raychaudhuri 1983<br />
Ghosh et al. 1971<br />
Patch 1939<br />
Raychaudhuri 1983<br />
Raychaudhuri 1983<br />
Raychaudhuri 1983<br />
Patch 1939<br />
Patch 1939<br />
(continued on next page)
14 Biological Control of Weeds: Southeast Asian Prospects<br />
Table 4.1.1 (continued)<br />
Species Location Other hosts References<br />
ALEYRODIDAE<br />
Bemisia tabaci<br />
DlASPlDlDAE<br />
Mycetaspis personata<br />
LYGAEIDAE<br />
Nysius inconspicuus<br />
Thysanoptera<br />
PHLAEOTHRIPIDAE<br />
Haplothrips gowdei<br />
THRlPlDAE<br />
Calipthrips ipomoeae<br />
Microcephalothrips<br />
abdominalis<br />
Thrips tabaci<br />
Diptera<br />
AGROMYZIDAE<br />
Calycomyza sp.<br />
Melanagromyza<br />
metallica<br />
CHLOROPIDAE<br />
Olcella pleuralis<br />
TEPHRITIDAE<br />
Xanthaciura imecta<br />
Lepidoptera<br />
ARCTll DAE<br />
Pareuchaetes<br />
pseudoinsulata<br />
(= Ammalo insulata)<br />
GELECHllDAE<br />
Dichomeris sp.<br />
NOCTUIDAE<br />
Pseudoplusia includem<br />
(=Plusia 00)<br />
Spodoptera frugiperda<br />
India, a very wide range Ang et al. 1977,<br />
Malaysia, Sastry 1984,<br />
Turkey Shreni et al. 1979<br />
Brazil polyphagous dlAraujo e Silva et al. 1968a<br />
India sesame and many Thangavelu 1978<br />
others<br />
Hawaii vector of pineapple Sakimura 1937<br />
yellow spot virus<br />
Brazil polyphagous dlAraujo e Silva et al. 1968a<br />
India polyphagous Gopinathan et al. 198 1<br />
Hawaii vector of pineapple Sakimura 1937<br />
yellow spot virus<br />
USA Spencer & Steyskal 1986<br />
India, etc no other host Singh & Beri 1973<br />
mentioned<br />
Trinidad C. odorata, C. ivaefolia, McFadyen 1988a<br />
C. iresinoides,<br />
Fleischmannia<br />
microstemon,<br />
Wedelia<br />
caracasana,<br />
Wulfla baccata<br />
Florida, C. odorata,<br />
Trinidad F. microsternon<br />
W. caracasana<br />
McFadyen 1988a,<br />
Needham 1946<br />
Nigeria, C. odorata Bennett & Cruttwell 1973,<br />
Trinidad Olaoye 1974<br />
Trinidad C. odorata Bennett & Cruttwell 1973<br />
Brazil polyphagous d'Araujo e Silva et al. 1968a<br />
Brazil polyphagous d'Araujo e Silva et al. 1968a<br />
(continued on next page)
MITES<br />
4.1 Ageratum conyzoides 15<br />
Species Location Other hosts References<br />
PYRALIDAE<br />
Pionea upalusalis Trinidad, C. odorata, C. ivaefolia, McFadyen 1988a<br />
Puerto Rico, Austroeupatorium<br />
Venezuela inulaefolium,<br />
Fleischmannia<br />
microstemon<br />
Brevipalpus obovatus India cotton, Solanum nigrum, Sadana et al. 1983<br />
Sonchus asper,<br />
Phaseolus vulgaris,<br />
Euphorbia hirta,<br />
Xanthium sp.,<br />
Cichorium intybus<br />
Tetranychus urticae China a very wide range Dong et al. 1986<br />
Table 4.1.2 Natural enemies of Ageratum conzoides: nematodes.<br />
Species Location Other hosts References<br />
Aphelenchoidesfiagariae<br />
Helicotylenchus multicinctus<br />
Meloidogyne sp.<br />
Meloidogyne arenaria<br />
Meloidogyne arenaria<br />
thamesis<br />
Meloidogyne incognita<br />
Meloidogyne javanica<br />
Pratylenchus pratensis<br />
Rotylenchulus reniformis<br />
Hawaii strawberry, Vanda<br />
orchids, Impatiens,<br />
Nephrolepis biserrata<br />
Brazil banana, Portulaca<br />
oleracea and several<br />
weeds<br />
Cuba Eleusine indica,<br />
Croton lobatus,<br />
Cynodon dactylon<br />
Philippines<br />
Philippines<br />
Philippines many vegetables<br />
and weeds<br />
Philippines, many vegetables<br />
Nigeria and weeds<br />
Hawaii<br />
Hawaii, India many weeds<br />
Sher 1954<br />
Zem & Lordello 1983<br />
Acosta et al. 1986<br />
Holm et al. 1977<br />
Valdez 1968<br />
Valdez 1968<br />
Mamaril& Alberto 1989<br />
Mamaril& Alberto 1989,<br />
-Salawu et al. 1991<br />
Valdez 1968<br />
Holm et al. 1977<br />
Linford & Yap 1940,<br />
La1 et al. 1978
Biological Control of Weeds: Southeast Asian Prospects<br />
Table 4.1.3 Natural enemies of Ageratum conyzoides: fungi, bacteria and viruses.<br />
Species Location Other hosts References<br />
FUNGI<br />
Cercospora agerati<br />
Colletotrichium sp.<br />
Cylindrocladium<br />
quinqueseptalum<br />
Mycovellosiella perfoliata<br />
Puccinia conoclinii<br />
Sclerotium rolfsii<br />
BACTERIA<br />
Pseudomonas<br />
solanacearum<br />
India<br />
India<br />
India<br />
India<br />
many commercial<br />
hosts<br />
many<br />
India potato, Ranunculus<br />
sceleratus<br />
Stevens 1925<br />
Kulkami & Sharma 1976<br />
Sulochana et al. 1982<br />
Srivastava 198 1<br />
Stevens 1925<br />
Desai et al. 1980<br />
Sathiarajan &<br />
Sasikumar 1977,<br />
Sunaina et al. 1989<br />
VIRUSES<br />
Ageratum vein yellowing India, (transmitted by<br />
Ang et al. 1977,<br />
Malaysia Bemisia tabaci)<br />
Shreni et al. 1979<br />
anemone mosaic<br />
Holm et al. 1977<br />
Bidens mottle several, including<br />
Zinnia, petunia<br />
& Verbena<br />
Logan & Zettler 1984<br />
hibiscus yellow vein India (transmitted by<br />
Jeyarajan et al. 1988<br />
mosaic<br />
B. tabaci)<br />
pineapple yellow spot Hawaii<br />
Sakimura 1937<br />
potato virus Y<br />
India<br />
potato<br />
Joshi & Prakash 1977<br />
tapioca mosaic<br />
India<br />
(transmitted by<br />
B. tabaci)<br />
Jeyarajan et al. 1988<br />
tobacco leaf curl India tomato<br />
Holm et al. 1977,<br />
Reddy et al. 1981<br />
tomato leaf curl Turkey, many weeds<br />
Sastry 1984,<br />
India<br />
(transmitted by<br />
B. tabaci)<br />
Jeyarajan et al. 1988<br />
urd bean yellow mosaic India (transmitted by<br />
B. tabaci)<br />
Jeyarajan et al. 1988<br />
Zinnia yellow net India (transmitted by<br />
B. tabaci)<br />
Srivastava et al. 1977
18 Biological Control of Weeds: Southeast Asian Prospects<br />
Amaranthus spinosus<br />
(after Holm et a/. 1977)
Map 4.2 Amaranthus spinosus<br />
4.2 Amaranthus spinosus 19<br />
Amaran thus spinosus<br />
Mass rearing and release, as required, of the weevil Hypolixus trunculatus is reported to<br />
provide good control of Amaranthus spinosus in Thailand but, of course, this is augmen-<br />
tative rather than classical biological control.<br />
Three other insects (a weevil, a leaf mining fly and a caterpillar) are known which may<br />
prove to be adequately specific for classical biological control.<br />
However, almost nothing is known about the natural enemies of A. spinosus in tropical<br />
America where it evolved and it would thus be necessary to cany out a survey there in<br />
order to evaluate what potential biological control agents are available.
20 Biological Control of Weeds: Southeast Asian Prospects<br />
4.2 Amaranthus spinosus L.<br />
Amaranthaceae<br />
spiny amaranth, spiny pigweed, needle burr; hin nu nive tsu bauk (Myanmar),<br />
phak khom nam (Thailand), phti banla (Cambodia), bayam duri (Malaysia and<br />
Indonesia), orai (Philippines), den gai (Vietnam)<br />
Rating<br />
+++ Myan, Thai, Phil<br />
17 ++ Msia, Sing<br />
+ Laos, Camb, Viet, Indo<br />
Origin<br />
Tropical America.<br />
Distribution<br />
A. spinosus is mainly tropical and subtropical in distribution, but also extends into the<br />
temperate zone from latitude 30°N to 30"s.<br />
Characteristics<br />
A. spinosus is an erect, much branched, annual, growing to 1.2 m. Its stems are angled in<br />
cross section, fleshy, often reddish and bear many spines. Its leaves are alternate, with a<br />
pair of straight spines up to 1 cm long at the base. The inflorescence is long, slender and<br />
terminal or arises from leaf axils. The flowers are small, greenish and unisex. It is propa-<br />
gated by reddish brown seeds.<br />
Importance<br />
Spiny amaranth prospers in warm sunny situations, but not where it is cool or shady. It is<br />
not reported as a problem in the Mediterranean or Middle East. It is a weed in 44 coun-<br />
tries in 28 crops, mainly in the Caribbean, in the west and south of Africa, in India and in<br />
Southeast Asia. Up to 235 000 seeds per plant have been recorded. Seeds are spread by<br />
wind and water. Some germinate soon, others over several months and still others remain<br />
viable in the soil for many years. A. spinosus is abundant in cultivated and abandoned<br />
fields, along roadsides and in waste places. It is a weed of varying degrees of aggressive-<br />
ness in many crops, including upland rice, cotton, cowpeas, groundnuts, maize, mangos,<br />
millet, pineapples, sugarcane and vegetables. The rigid needle-like spines break off in the<br />
hands of workers in sugarcane, cotton and other crops.<br />
A. spinosus may contain high nitrate levels and has been implicated in livestock poi-<br />
soning. It is avoided by most animals because of its spines. Leaves are sometimes used<br />
by humans as a green vegetable. Other Amaranthus species are valuable as a grain crop<br />
in some South American countries and the family Amaranthaceae contains a number of<br />
widely grown ornamental garden species (Purseglove 1968).
4.2 Amaranthus spinosus 21<br />
Natural enemies<br />
A. spinosus is attacked by a number of natural enemies (Tables 4.2.1 and 4.2.2), but most<br />
of the reports come from outside its native range and are of non-specific organisms. The<br />
agromyzid fly Haplopeodes minutus, known in USA from species of Amaranthus and<br />
Chenopodium (Spencer and Steyskal 1986) and both the beetle Cassida nigriventris and<br />
the moth Coleophora versurella, known in Pakistan from these same plant genera (Khan<br />
et al. 1978), may prove to be sufficiently specific to be candidate biological control<br />
agents.<br />
The weevil Hypolixus trunculatus, whose larvae tunnel in the stems and form galls,<br />
is known from Pakistan, India and Thailand and attacks Amaranthus spinosus, A. viridis<br />
and Digera arvensis. Although it has a relatively long life'cycle and low reproductive<br />
capacity, mass rearing and augmentative releases have resulted in a satisfactory level of<br />
control and replaced the use of herbicides in Thailand (Julien 1992, Napompeth 1982,<br />
1989, 1992a). Females deposit eggs singly in cavities scooped out of the shoots. Larvae<br />
tunnel down inside the stem to its base, where a gall develops. Breeding continues<br />
throughout the year but is at its height in late summer. At this time the life cycle is 44 to<br />
50 days. Pupation occurs within the gall. Larvae and pupae are parasitised by larvae of<br />
the pteromalid wasp Oxysychus sp. (Aganval 1985).<br />
Evans (1987) records five fungi from A. spinosus but, except for one which is<br />
unsuitable because it has a wide host range, too little is known about their host specificity<br />
to assess the prospects for their use in classical biological control.<br />
Comment<br />
Almost nothing is known about the natural enemies of A. spinosus in tropical America<br />
where it evolved. A survey in this region would be necessary to document the organisms<br />
attacking it. There are good general grounds for believing that there are some natural<br />
enemies that are specific to the family Amaranthaceae. In most countries, members of<br />
this family have little value as crop plants, so the chances are that some safe natural ene-<br />
mies will be found that are of value as classical biological control agents.<br />
Table 4.2.1 Natural enemies of Amaranthus spinosus: insects and mites.<br />
Species Location Other hosts References<br />
INSECTS<br />
Hemiptera<br />
APHlDlDAE<br />
Myzus persicae Malawi, highly polyphagous Chapola 1980, Napompeth<br />
Thailand 1982<br />
COREIDAE<br />
CIetus fuscescens Nigeria Amaranthus dubius, Ukwela & Ewete 1989<br />
A. cruenrus,<br />
A. hypochondriachus<br />
LYGAEIDAE<br />
Germalus unipunctarus Vanuatu Cock 19841,<br />
Nysius sp. Vanuatu Cock 1984b<br />
(continued on next page)
22 Biological Control of Weeds: Southeast Asian Prospects<br />
Table 4.2.1 continued<br />
Species Location Other hosts References<br />
MlRlDAE<br />
Horcias nobilellus Brazil ' polyphagous d'Araujo e Silva et al. 1968a<br />
PI ESMATIDAE<br />
Piesma cinereum Brazil polyphagous d'Araujo e Silva et al. 1968a<br />
Thysanoptera<br />
PHLAEOTHRIPIDAE<br />
Haplothrips India Amaranthus viridis, Dhiman 1986<br />
longisetosus A. oleosa, Chenopodium<br />
anthelminthicum<br />
Coleoptera<br />
CHRYSOMELIDAE<br />
Cassida exilis<br />
Cassida nigriventris<br />
CURCULIONIDAE<br />
Ceutorhynchus<br />
asperulus<br />
Hypolixus<br />
trunculatus<br />
MELYRIDAE<br />
Astylus lineatus<br />
Pakistan Amaranthus viridis,<br />
Chenopodium album<br />
Pakistan Amaranthus viridis,<br />
Chenopodium album,<br />
Spinucia oleracea<br />
India red gram, Amaranthus<br />
viridis, A. tricolor,<br />
Basella alba<br />
Pakistan, Amaranthus viridis,<br />
India, Chromolaena odorata<br />
Thailand Digera arvensis<br />
Brazil citrus<br />
Baloch et al. 1976<br />
Baloch et al. 1976<br />
Khan et al. 1978<br />
Puttaswamy &<br />
Channabasavannal98 1,<br />
Puttaswamy et al. 198 1<br />
Aganval 1985,<br />
Baloch et al. 1976, 1977,<br />
Ghani 1965, Julien 1992<br />
Napompeth 1982, 1990b,<br />
1992a<br />
d'Araujo e Silva et al. 1968a<br />
Diptera<br />
AGROMYZIDAE<br />
Haplopeodes USA Amaranthus, Spencer & Steyskal 1986<br />
minutus Chenopodium<br />
Lepidoptera<br />
COLEOPHORIDAE<br />
Coleophora<br />
versurella<br />
CURCULIONIDAE<br />
Hypolixus ritsemae<br />
LYCAENIDAE<br />
Zizeeria knysna<br />
Zizeeria krupta<br />
NOCTUIDAE<br />
Neogalea<br />
(= Spodoptera) sunia<br />
Spodoptera eridania<br />
Spodoptera exigua<br />
. Spodoptera litura<br />
1<br />
Pakistan<br />
Vanuatu<br />
Pakistan<br />
Pakistan<br />
Nicaragua<br />
Nicaragua<br />
Nicaragua<br />
Philippines<br />
Chenopodium botrys Khan et al. 1978<br />
Cock 19841,<br />
Baloch et al. 1976<br />
Baloch et al. 1977, Ghani<br />
1965<br />
polyphagous Savoie 1988<br />
polyphagous Savoie 1988<br />
polyphagous Savoie 1988<br />
highly polyphagous Moody et al. 1987<br />
(continued on next page)
4.2 Amaranthus spinosus 23<br />
Species Location Other hosts References<br />
PYRALI DAE<br />
Loxostege sp. Argentina seed heads of<br />
Amaranthus sp.<br />
(the genus Loxostege<br />
Spoladea (=Hymenia)<br />
recurvalis<br />
SCYTHRIDIDAE<br />
Eretmocera<br />
impactella<br />
TORTRICIDAE<br />
Archips sp.<br />
YPONOMEUTIDAE<br />
Plutella xylostella<br />
MITE<br />
TETRANYCHIDAE<br />
Tetranychus<br />
novocaledonicus<br />
India,<br />
Pakistan<br />
Vanuatu<br />
contains pests)<br />
polyphagous<br />
Pakistan Amaranthus viridis,<br />
Chenopodium album<br />
Pakistan<br />
Pakistan<br />
India Amaranthus tricolor,<br />
4. viridis<br />
C.J. Deloach<br />
pers. comm. 1980<br />
Baloch et al. 1976<br />
Chaudhury & Kapil 1977,<br />
Lock 1984b. Ghani 1965<br />
Baloch et al. 1977<br />
Ghani 1965<br />
Ghani 1965<br />
Puttaswav.y dr<br />
Channabasavanna 1981<br />
Table 4.2.2 Natural enemies of Amaranthus spinosus: nematodes, fungi, viruses.<br />
Species Location Other hosts References<br />
NEMATODES<br />
Cactodera amaranthi Cuba spinach, other species Stoyanov 1972<br />
Meloidogyne incognita<br />
Pratylenchus zeae<br />
Pseudocephalobus indicus<br />
Philippines<br />
of Amaranthus<br />
rice, many weeds<br />
India only recorded on<br />
Valdez 1968<br />
Fortuner 1976<br />
Joshi 1972<br />
Rotylenchulus reniformis India, USA<br />
A. spinosus<br />
many weed hosts Inserra et al. 1989,<br />
La1 et al. 1978<br />
FUNGI<br />
Albugo bliti Dominica, many Amaranthaceae<br />
Jamaica, India,<br />
Pakistan, Sudan<br />
Alternaria compacta<br />
Aposphaeria amaranthi<br />
Bipolaris indica (as<br />
Drechslera indica)<br />
India<br />
USA potential bioherbicide<br />
for A. albus; effect on<br />
A. spinosus not known<br />
many, including<br />
Helianthus, Pennisetum,<br />
Portulaca<br />
Baloch et al. 1977,<br />
Evans 1987<br />
Kar & Ashok-Das 1988<br />
Mintz & Weidemann 1992<br />
Evans 1987,<br />
Kenfield et al.<br />
1989<br />
(continued on next page)
24 Biological Control of Weeds: Southeast Asian Prospects<br />
Table 4.2.2 continued<br />
Species Location Other hosts References<br />
Cercospora brachiata<br />
(= C. amaranthi)<br />
Fusarium oxysporum f.sp.<br />
elaeidis<br />
Phoma tropica<br />
Puccinia sp.<br />
VIRUSES<br />
cucumber mosaic<br />
Digera mosaic<br />
groundnut rosette<br />
tobacco bunchy top<br />
tobacco mosaic<br />
India, Nigeria, many Amaranthaceae<br />
Uganda, Trinidad,<br />
USA, Japan,<br />
China, USSR<br />
Nigeria oil palm, Chromolaem<br />
odorata, Impera ta<br />
cylindrica, Mariscus<br />
alternifolius<br />
India<br />
Hong Kong<br />
India cucumber, Solanum,<br />
nigrum, Tagetes<br />
erecta, etc<br />
India several weeds<br />
Malawi (Myzus persicae<br />
Philippines is a vector)<br />
Evans 1987<br />
Oritsejafor 1986<br />
Evans 1987<br />
Evans 1987<br />
Suteri et al. 1980<br />
Singh et al. 1975<br />
Adams 1967<br />
Chapola 1980<br />
Eugenio & del Rosario 1962
Biological Control of Weeds: Southeast Asian Prospects<br />
Bidens pilosa<br />
(after Holm et a/. 1977)
Map 4.3 Bidens pilosa<br />
4.3 Bidens pilosa<br />
Bidens pilosa is native to tropical America. Preliminary studies, based mainly on<br />
Trinidad, indicate that it is attacked by a number of natural enemies, mainly insects, and<br />
that several of these may be sufficiently host specific to be considered as biological con-<br />
trol agents. Further host specificity studies are required and additional, wider-ranging<br />
searches, particularly in South America.
28 Biological Control of Weeds: Southeast Asian Prospects<br />
4.3 Bidens pilosa L.<br />
Asteraceae<br />
cobbler's pegs, Spanish needle; djaringan ketul (Indonesia), pisau pisau<br />
(Philippines) yah koen jam khao (Thailand)<br />
Rating<br />
++ Thai, Indo, Phil<br />
10 + Myan, Laos, Camb, Viet<br />
Msia<br />
Origin<br />
Tropical America<br />
Distribution<br />
Pantropical. Known from Java before 1835, but apparently not present in Kalimantan or<br />
the Moluccas (Soerjani et al. 1987).<br />
Characteristics<br />
Bidens pilosa is an erect, slender, branching, annual herb growing up to 1.5 m. Its stems<br />
are four-angled in cross section and its leaves opposite and sparsely hairy. The abundant<br />
yellow flowers are borne in heads on long stalks and produce black, barbed seeds charac-<br />
teristically radiating in all directions from a common base. The recurved, 2-toothed barbs<br />
enable the seeds to stick readily to hair and clothing and they are also distributed by wind<br />
and water. Cobbler's pegs prefers moister soils and flowers all year round.<br />
Importance<br />
A very common weed of 31 crops in more than 40 countries, B. pilosa occurs in gardens,<br />
cultivated land, open waste places and along roadsides. It is an important weed of pas-<br />
tures, maize, sorghum, vegetables, cotton, tea, coffee, cassava, coconut, oil palm, citrus,<br />
papaya, rice, rubber and tobacco. Single plants produce up to 6000 seeds, many of which<br />
germinate readily, permitting three or four generations a year in some regions.<br />
Some seeds remain viable in the soil for at least 5 years. When herbicides have<br />
eradicated perennial grasses this weed often becomes dominant.<br />
In South Africa the early spring growth is sometimes eaten by humans, but has low<br />
nutritive value. It has a pungent essential oil that may taint milk.<br />
Natural enemies<br />
These 'are also dealt with in 'Biological Control: Pacific Prospects' (Waterhouse and<br />
Norris 1987) which did not assess B. pilosa a particularly promising target for biological<br />
control. However, more information has since become available (Table 4.3.1 and 4.3.2),
4.3 Bidens pilosa 29<br />
particularly concerning leaf miners and seed head feeders of the fly family Agromyzidae.<br />
This suggests that there may be good prospects for some of these natural enemies.<br />
Few details are available of the natural enemies of B. pilosa in Brazil. The pupal<br />
stage of the chrysomelid beetle Phaedon pertinax (= P. consimilis) lasts 6 to 8 days and<br />
the pentatomid bug Stiretrus erythrocephalus passes through 4 instars in 30 days<br />
(Ribeiro 1953). Thrips killed 22.25% of B. pilosa plants (particularly seedlings) and<br />
Diptera infested 97.8% of flower heads. Parasitisation of these Diptera by wasps and<br />
flies, varied from 40.96% to 58.91 % according to the size of the population (Esposito et<br />
al. 1985).<br />
About half of the 2500 species of the family Agromyzidae have known hosts and<br />
almost all of this group are restricted in their feeding to a single family or genus. Only 16<br />
of the species (0.6% of the total) are truly polyphagous, feeding on a number of unrelated<br />
families (Spencer 1990). Agromyzid flies are, therefore, worth serious consideration as<br />
classical biological control agents. In this context, plants of the genus Bidens appear to be<br />
particularly attractive to agromyzid flies for they support 19 species (Table 4.3.3).<br />
In the tribe Coreopsideae (of the family Asteraceae) only two (Bidens and<br />
Coreopsis) of its 26 genera support Agromyzidae (Table 4.3.4). Coreopsis is native in<br />
North America, but no agromyzids are known on it there, although three polyphagous<br />
species are known to attack it in Europe, India or Australia (Spencer 1990).<br />
Eleven of the above 19 species are known from Bidens pilosa (Table 4.3.1). Of<br />
these, three are restricted to the genus Bidens (perhaps even to B. pilosa), two are<br />
polyphagous, and the remaining six have one or more additional hosts in other genera of<br />
the Asteraceae. Ten of the eleven species are restricted to the Americas and further host<br />
specificity tests may well indicate that many are valuable biological control agents. Four<br />
of the ten form blotch mines (Arnauromyza maculosa, Calycomyza allecta, C. platyptera<br />
and Liriomyza archboldi), one makes long, linear irregular mines (Liriomyza venegasiae),<br />
and three feed in the seed heads (Liriomyza insignis, Melanagromyza bidentis and M.<br />
floris) (Spencer 1990, Spencer and Steyskal 1986).<br />
The flower heads of B. pilosa are also attacked by three species of Tephritidae in<br />
Central America and by one of these in India. Adult weevils of the genera Baris,<br />
Centrinaspis and Promecops feed in the flowers of B. pilosa and other Asteraceae, but<br />
are thought not to breed there. Several other insects (at least three other beetles and a<br />
pierid butterfly) have also been recorded from B. pilosa and sometimes from other<br />
Asteraceae as well.<br />
Table 4.3.4 shows the position of the genus Bidens as a member of the tribe<br />
Coreopsidae, within the family Asteraceae. There may well be natural enemies that<br />
attack it, but not any species of agricultural or special environmental significance.<br />
Attempts at biological control<br />
There have been none.
30 Biological Control of Weeds: Southeast Asian Prospects<br />
Table 4.3.1 Natural enemies of Bidens pilosa: insects.<br />
Species Location Other hosts References<br />
Hemiptera<br />
ALEYRODI DAE<br />
Dialeurodes vulgaris India coffee, Erythrina<br />
lithosperma<br />
Venkataramaiah 1974<br />
APHlDlDAE<br />
Aphis coreopsidis Brazil soybean Almeida 1979, d'Araujo<br />
e Silva et al. 1968a<br />
d'Araujo e Silva et al. 1968a<br />
Christie et al. 1974,<br />
d'Araujo e Silva et al. 1968a<br />
Aphis illinoisensis Brazil<br />
Uroleucon<br />
(= Dacrynotus) sp.<br />
MlRlDAE<br />
Brazil, USA tobacco, lettuce<br />
Garcanus gracilentus Brazil sweet potato,<br />
polyphagous<br />
d'Araujo e Silva et al. 1968a<br />
Horcias nobilellus<br />
PENTATOMIDAE<br />
Brazil polyphagous<br />
Amaranthus spinosus<br />
d'Araujo e Silva et al. 1968a<br />
Stiretrus erythrocephalus Brazil<br />
Ribeiro 1953<br />
Thyanta perditor Brazil soybean<br />
Grazia et al. 1982<br />
Coleoptera<br />
APlONlDAE<br />
Apion luteirostre<br />
CHRYSOMELIDAE<br />
Chalcophana viridipennis<br />
Chlamisus insularis<br />
Phaedon pertinax<br />
(= P. consimilis)<br />
Physimerus pygmaeus<br />
CURCULIONIDAE<br />
Baris sp.<br />
South America Mikania micrantha Cock 1980<br />
Brazil<br />
Trinidad Chromolaena odorata,<br />
C. ivaefolia<br />
Brazil, Mikania micrantha<br />
(not in Trinidad)<br />
South America Mikania micrantha<br />
d'Araujo e Silva et al. 1968a<br />
McFadyen 1988a<br />
Cock 1980, d'Araujo e Silva<br />
et a]. 1968a, Ribeiro 1953<br />
Cock 1980<br />
Trinidad (feed in B. pilosa<br />
flowers)<br />
Cruttwell 1971a<br />
Centrinaspis sp. Trinidad (feed in B. pilosa<br />
flowers)<br />
Cruttwell 197 1 a<br />
Promecops sp. Trinidad (feed in B. pilosa<br />
flowers)<br />
Cruttwell 197 la<br />
Rhodobaenus<br />
Trinidad adults feed on stems, McFadyen 1988a<br />
cariniventris<br />
and petioles of B. pilosa,<br />
Chromolaena odorata,<br />
C. ivaefolia,<br />
Austroeupatorium<br />
inulaefolium<br />
Rhodobaenus,<br />
Trinidad feed in B. pilosa stems: McFadyen 1988a<br />
tredecimpunctatus<br />
and in several other<br />
Asteraceae<br />
Diptera<br />
AGROMYZIDAE<br />
Amauromyza maculosa Trinidad (also polyphagous, but favours<br />
N&S America, Asteraceae<br />
Hawaii)<br />
Cruttwell 197 1 a,<br />
Spencer 1990, Spencer &<br />
Steyskal 1986<br />
(continued on next page)
4.3 Bidens pilosa 31<br />
Species Location Other hosts References<br />
Calycomyza allecta Trinidad (also<br />
Brazil,<br />
Guadeloupe,<br />
Venezuela)<br />
Calycomyza USA (Florida,<br />
platyptera California)<br />
Liriomyza archboldi Florida<br />
(Bahamas,<br />
Costa Rica)<br />
Liriomyza insignis Costa Rica<br />
Liriomyza trifolii cosmopolitan<br />
Liriomyza venegasiae Southern<br />
California<br />
Liriomyza sp. Argentina<br />
Melanagromyza bidentis Florida,<br />
Caribbean<br />
Melanagromyzafloris Costa Rica,<br />
Mexico,<br />
Trinidad<br />
(also Florida,<br />
Neotropics)<br />
Melanagromyza splendid USA, Hawaii<br />
Phytomyza atricornis Australia<br />
CEClDOMYllDAE<br />
Asphondylia bidens Florida<br />
DROSOPHILIDAE<br />
Cladochaeta nebulosa Florida<br />
TEPHRlTlDAE<br />
Dioxyna sororcula Florida,<br />
(= D. picciola) Trinidad,<br />
widespread<br />
Xanthaciura insecta Florida,<br />
Trinidad<br />
Lepidoptera<br />
ARCTllDAE<br />
Hypercompe Brazil<br />
(=Ecpantheria)<br />
hambletoni<br />
NOCTUIDAE<br />
Cropia (=Dyops) minthe Brazil<br />
Mocis latipes Brazil<br />
Thysanoplusia Kenya<br />
(- Diachrysia) orichalcea<br />
PlERlDAE<br />
Perrhybris phaloe Trinidad<br />
(= Ascia buniae phaloe)<br />
Helianthus, Rudbeckia<br />
and garden Asteraceae<br />
Asteraceae, including<br />
Aster, Helianthus,<br />
Zinnia<br />
restricted to Bidens<br />
restricted to Bidens<br />
polyphagous, including<br />
Chrysanthemum<br />
Venegasia carpesioides<br />
restricted to Bidens<br />
Verbesina sp.<br />
Calendula sp<br />
Asteraceae including<br />
Helianthus, Lactuca<br />
polyphagous, including<br />
Cineraria<br />
attacks several<br />
Asteraceae in India<br />
Ageratum conyzoides,<br />
Chromolaena odorata<br />
Fleischmannia<br />
caracasana<br />
Panicum maximum,<br />
Paspalum notarum,<br />
Hyparrhenia rufa<br />
coffee and other crops<br />
Cruttwell 1971b, Frick<br />
19.56, Spencer 1990,<br />
Spencer & Steyskal 1986<br />
Spencer 1990<br />
Spencer & Steyskal 1986<br />
Spencer 1990,<br />
Spencer & Steyskal 1986<br />
Spencer 1990<br />
Spencer 1990,<br />
Spencer & Steyskal 1986<br />
Spencer 1990<br />
Spencer 1990<br />
Cruttwell 197 la, Spencer<br />
1990, Spencer &<br />
Steyskal 1986<br />
Spencer 1990,<br />
Spencer & Steyskal 1986<br />
Kleinschmidt 1970<br />
Steyskal 1972<br />
Steyskal 1972<br />
Cruttwell 1971 a, 1972a,b,<br />
Steyskal 1972<br />
McFadyen 1988a,<br />
Steyskal 1972<br />
d'Araujo e Silva et al. 1968a<br />
d'Araujo e Silva et al. 1968a<br />
Lourencao et al. 1982<br />
Bardner & Mathenge 1974<br />
Cruttwell 197 1 a
32<br />
Biological Control of Weeds: Southeast Asian Prospects<br />
Table 4.3.2 Natural enemies of Bidens pilosa: nematodes, fungi, mycoplasmas,<br />
viruses.<br />
Species Location<br />
NEMATODES<br />
Meloidogyne sp. Hawaii<br />
Meloidogyne hapla India<br />
Pratylenchus minutus Hawaii<br />
Rotylenchulus reniformis USA<br />
FUNGI<br />
Cercospora bidenfis Mauritius<br />
Cercospora rnegalopotamica Hawaii<br />
Entyloma guaraniticurn Mauritius<br />
Uromyces bidenticola Hawaii, Mauritius<br />
MYCOPLASMAS<br />
aster yellows Hawaii<br />
Bidens witches broom Brazil<br />
VIRUSES<br />
Bidens mosaic<br />
groundnut rosette<br />
Sonchus yellow net<br />
soybean mosaic<br />
tomato spotted wilt<br />
Brazil<br />
Hawaii<br />
Florida<br />
Brazil<br />
Hawaii<br />
PARASITIC PLANT<br />
Cassytha filiformis Hawaii<br />
Table 4.3.3 Species in Agromyzid genera attacking Bidens.<br />
References<br />
Linford et al. 1949<br />
Singh et al. 1979<br />
Linford et al. 1949<br />
Inserra et al. 1989, McSorley<br />
et al. 198 1<br />
Rochecouste & Vaughan 1959<br />
Stevens 1925<br />
Rochecouste & Vaughan 1959<br />
Anon 1960, Rochecouste &<br />
Vaughan 1959<br />
Holm et al. 1977<br />
Vega et al. 198 1<br />
Kuhn et al. 1982<br />
Adams 1967<br />
Christie et al. 1974<br />
Almeida 1979<br />
Sakimura 1937<br />
Raabe 1965<br />
Genus Specific to Bidens Specific to Coreopsideae Polyphagous<br />
Melanagromyza<br />
Amauromyza<br />
Liriomyza 3<br />
Calycomyza<br />
Chromatomyia<br />
Total 3
4.3 Bidens pilosa 33<br />
Table 4.3.4 Relationship of four major Southeast Asian weeds and some economically<br />
important genera within the family Asteraceae.<br />
Family Asteraceae: 21 000 species (Mabberley 1987)<br />
Tribe Some economically Weed species<br />
important genera<br />
Arctoteae<br />
Carlineae<br />
Echinopsideae<br />
Cardueae<br />
Mutisieae<br />
Lactuceae<br />
Vemonieae<br />
Inuleae<br />
Astereae<br />
Eupatorieae<br />
Calenduleae<br />
Senecioneae<br />
Anthemideae<br />
Heleniae<br />
Madieae<br />
Heliantheae<br />
Tageteae<br />
Coreopsideae<br />
Carthamnus, Cynara<br />
Cichorium, Lactuca<br />
Aster<br />
Cineraria<br />
Chrysanthemum<br />
Dahlia<br />
Cosmos, Helianthus, Zinnia<br />
Ageratum conyzoides,<br />
Chromolaena odorata,<br />
Mikania micrantha<br />
Coreopsis Bidens pilosa<br />
The family Asteraceae, by far the largest in the dicotyledons, has been subdivided into<br />
18 tribes, some 1300 genera and about 21 000 species (Mabberley 1987). It contains sur-<br />
prisingly few economically important crop plants, of which lettuce (Lactuca sativa),<br />
sunflower (Helianthus annuus) and globe artichoke (Cynara scolymus) are the major<br />
species. However, there are a number of commercially important garden plants, espe-<br />
cially in the genus Chrysanthemum.
Biological Control of Weeds: Southeast Asian Prospects<br />
Chromolaena odorata<br />
(after Holm etal. 1977)
Map 4.4 Chromolaena odorata<br />
4.4 Chromolaena odorata 35<br />
Chromolaena odorata<br />
Chromolaena odorata is not a problem weed in the tropical Americas where it evolved. It<br />
is attacked there by more than 200 insects, at least a quarter of which are probably suffi-<br />
ciently host specific to be considered as classical biological control agents. The aggres-<br />
siveness of C. odorata in countries to which it has spread is probably due to the absence<br />
of most of these natural enemies.<br />
The arctiid moth Pareuchaetes pseudoinsulata has been established in India,<br />
Sri Lanka, Philippines, Sabah (Malaysia), the Mariana Is (Guam, Rota, Saipan, Tinian,<br />
Aguijan) and Federated States of Micronesia (Yap, Pohnpei, Kosrae), but only in the two<br />
latter island groups has it had spectacular success in controlling the weed. The mite<br />
Acalitus adoratus has spread naturally to Southeast Asia and Micronesia but, as yet, is<br />
having minor impact.<br />
It is probable that a group of natural enemies will be necessary to bring about effec-<br />
tive biological control of C. odorata in Southeast Asia, but there are a number of species<br />
that are well worthy of attention and longer term prospects for control appear promising.
36<br />
Biological Control of Weeds: Southeast Asian Prospects<br />
4.4 Chromolaena odorata (L.) R.M. King and H. Robinson<br />
(Formerly Eupatoriurn odoraturn)<br />
Asteraceae<br />
Siam weed, devil weed; bizat, tawbizat (Myanmar), tontrem khet (Cambodia),<br />
French weed (Laos), pokok tjerman (Malaysia), kirinyu, kumpai jepang, rumput<br />
go1 kar (Indonesia), hagonoy (Philippines) saab sua, yah sua mop (Thailand), co<br />
hoi (Vietnam)<br />
Rating<br />
+++ Msia, Phil<br />
18 ++ Myan, Thai, Laos, Camb, Viet, Indo<br />
Origin<br />
Central America and tropical South America (from Florida to northern Argentina).<br />
Distribution<br />
C. odorata is a weed throughout Southeast Asia, Irian Jaya, Papua New Guinea, New<br />
Britain, Mariana and Caroline Is, southern China, Taiwan, Sri Lanka, Bangladesh, India,<br />
West, Central and South Africa.<br />
Characteristics<br />
C. odorata is an upright or scrambling, thicket-forming, perennial shrub, growing from<br />
1.5 to 3 m high. Its roots are fibrous with a few well formed anchor roots and many later-<br />
als, the stems round, yellowish, hairy or almost smooth and profusely branched. Its<br />
leaves are opposite, with toothed margins and are conspicuously three veined. The flow-<br />
ers are at the tips of all stems, in clusters of 20 to 60, white or pale lilac. The achenes<br />
consist of 5 mm-long seeds with hooks on their angles, together with a pappus of 5 mm-<br />
long white bristles. The leaves have a pungent odour when damaged. Seed production is<br />
prolific (as many as 2 million per plant) and seeds provide the main mode of reproduc-<br />
tion. The achenes float long distances in the air and the seed hooks cling to hair and<br />
clothing. Germination occurs as soon as there is adequate moisture, although some 66%<br />
of seeds are not viable. Buried seeds lose up to 50% of their viability after 2 years (Yadav<br />
and Tripathi 1982).<br />
Importance<br />
C. odorata is not a serious weed in the Americas and no specific control methods are<br />
necessary (McFadyen 1991a). This is in stark contrast to its serious weed status in the<br />
countries to which it has spread and has been attributed to the many natural enemies that<br />
attack it in the Americas (McFadyen 1989, 1991~). It was introduced to Calcutta in the<br />
1840s, had spread into Sri Lanka, Southeast Asia and Nigeria by the 1940s and into Irian<br />
Jaya, New Britain and Micronesia by the 1980s. It is forecast to spread widely and
4.4 Chromolaena odorata 37<br />
aggressively in equatorial Africa, northern and eastern Australia and the Pacific<br />
(McFadyen 1988b, 1989).<br />
C. odorata grows in many soil types, but prefers well drained conditions and an<br />
annual rainfall above 1000 mm. Although it is not a problem in continuously cultivated<br />
land, it is most common and causes most losses in plantation crops, including coconut,<br />
rubber, oil palm, tea, coffee, cocoa, teak and cashew. It also thrives in areas newly<br />
cleared for planting, in abandoned or neglected fields, wastelands and along roadsides. It<br />
is sometimes a weed in pastures. Its rapid growth enables it to smother most competitors<br />
and it inhibits many with its allelopathic properties. It dies back after flowering in areas<br />
with a pronounced dry season and then becomes a fire hazard. After burning or cutting,<br />
the plants shoot freely from the crown. They are capable of forming dense tangled bushes<br />
two to three metres high, occasionally reaching six metres as climbers on other plants.<br />
The stems branch freely, with 20 or more laterals developing from axillary buds and<br />
often bent over under their own weight. Impenetrable stands of the weed cut off access to<br />
pastures and provide hiding places for rats, pigs and other undesirable animals. C.<br />
odorata is intolerant of shade, so that it dies out when the canopy closes in plantations<br />
(Ambika and Jayachandra 1990, McFadyen 1988b, 199 1 a). The shoots and young leaves<br />
contain nitrate at levels 5 to 6 times those toxic to stock and also pyrrolizidine alkaloids<br />
and cattle deaths occur following grazing. Hand weeding of Chromolaena is reported to<br />
cause skin allergy and scratches to result in infections (Ambika and Jayachandra 1990).<br />
It is interesting that the spread of C. odorata in West Africa has led to a<br />
polyphagous grasshopper Zonocerus variegatus becoming a pest. Although they are<br />
unable to mature on the weed as the only diet, hoppers are strongly attracted to the plant<br />
and especially to its flowers; and thickets are preferred night roosting sites. BopprC<br />
(1 991) hypothesises that the pyrrolizidine alkaloids accumulated from feeding on C.<br />
odorata protect the grasshoppers and their eggs from predators and parasitoids, leading<br />
to increased fitness and population density. However, this only occurs during the dry season,<br />
but not in the wet season when C. odorata does not bloom.<br />
Claims have been made (e.g. Field 1991, Herren-Gemmill 1991) that, under some<br />
circumstances, C. odorata may be beneficial to resource-poor farmers. One potential<br />
advantage, is its ability to outcompete another serious weed, alang-alang (Imperata cylindric~).<br />
However, McFadyen (1992) pointed out that a suitable perennial legume would<br />
be even more beneficial than C. odorata, and she also refuted a number of other claims.<br />
In Sri Lanka the indigenous legume Tephrosia purpurea has been successfully used to<br />
suppress weeds including C. odorata under coconut (Salgado 1972). Whatever potential<br />
benefits there may be in the presence of C. odorata there is an enormous body of fact to<br />
demonstrate that C. odorata has serious adverse effects on agricultural productivity in<br />
countries to which it has been introduced.<br />
Natural enemies<br />
A good deal is known about the insects attacking Chromolaena odorata, mainly as a<br />
result of studies aimed at biological control which started in the late sixties at the<br />
Commonwealth Institute of Biological Control Station in Trinidad. A number of scientists
38<br />
Biological Control of Weeds: Southeast Asian Prospects<br />
were involved, but principally R.E. McFadyen (nee Cruttwell) (Bennett and Cruttwell<br />
1973, Bennett and Rao 1968, Bennett and Yaseen 1975, Cock 1984a, Cock and Holloway<br />
1982, Cruttwell 1973a,b, 1974, 1977a,b, Cruttwell and Bennett 1969, McFadyen<br />
1988a,b, 199 1 a,b, Yaseen and Bennett 1977).<br />
An extensive bibliography dealing with all aspects of C. odorata, including its natural<br />
enemies and biological control, was compiled by Muniappan et al. (1988a), later<br />
supplemented in Chromolaena odorata Newsletters 3 (1990) and 6 (1992). The proceedings<br />
of three International Workshops on Biological Control of Chromolaena odorata,<br />
held in 1988, 1991 and 1993 also contain a wealth of up-to-date information.<br />
In the Americas C. odorata is attacked by at least 207 insect and 2 mite species<br />
(McFadyen 1988a). Of these, about half are probably polyphagous, a quarter are restricted<br />
to the Asteraceae and a quarter specific to Chromolaena. All stages of growth of the<br />
above ground parts of the plants are attacked, but the roots have not been examined<br />
(McFadyen 1991a) and not all regions where C. odorata occurs naturally were visited.<br />
For other regions of the world McFadyen (1988a) quotes records of 42 insect and 9 mite<br />
species, the vast majority of which are, or are likely to prove, polyphagous. Since then a<br />
few additional species have been recorded, all but one of which (an eriophyid mite, see<br />
India below) are likely to be polyphagous.<br />
In Trinidad, the cumulative effect of the natural enemies is great, between 25 and<br />
50% of all growing tips being destroyed. Seed germination is as low as 17% and many<br />
flowerheads fail to produce seed. Seedlings often succumb to the attack of stem and tip<br />
feeding insects and competitiveness and growth of established plants is greatly reduced<br />
by insect attack. At different sites and in different seasons damage is caused by different<br />
insects and, in general, is heaviest in shaded sites. Some of the insects are heavily<br />
attacked by parasitoids and if introduced without these to another country might prove to<br />
be even more effective.<br />
In addition to an arctiid moth (Pareuchaetes pseudoinsulata) and a weevil (Apion<br />
brunneonigrum), which have already had considerable attention paid to them (see next<br />
section), McFadyen (1 99 1 c) has nominated an additional 1 1 insects for priority evaluation<br />
(Table 4.4.1). Furthermore, others (Cruttwell 1974, Cock 1984a, Cock and Holloway<br />
1982, McFadyen 1988c, Muniappan and Viraktamath 1986) have suggested an additional<br />
22 species (Table 4.4.2) which were evidently considered less important by McFadyen<br />
(1991~). It is clear therefore that, if required, there are many promising candidates for<br />
detailed consideration. The additional species of Pareuchaetes suggested by Cock and<br />
Holloway (1982) have not been investigated in detail, but all are believed to breed on C.<br />
odorata or related species and several may be better adapted climatically and biologically<br />
than P. pseudoinsulata to conditions in many overseas countries.<br />
Although no special search has been carried out except in Trinidad and Tobago a<br />
number of fungal pathogens occurring on C. odorata are shown in table 4.4.3. Half of the<br />
records 'come from outside its area of origin and must, therefore, be suspected of having a<br />
wider than desirable host range. Possibly Cionothrix praelonga is of greatest interest,<br />
since preliminary tests indicate that it may be host specific (Ooi et al. 1991). It is autoecious<br />
(i.e. it does not have an alternate host), occurs in the Caribbean and Venezuela and
4.4 Chromolaena odorata 39<br />
Table 4.4.1 Potential biological control agents for C. odorata: insects (after<br />
McFadyen 199 1 c).<br />
Species Part Damage Problem Country found<br />
attacked<br />
Coleoptera<br />
CHRYSOMELIDAE<br />
Aulacochlamys sp.<br />
Chlamisus insularis<br />
Pentispa explanata<br />
CURCULIONIDAE<br />
Rhodobaenus<br />
cariniventris<br />
Diptera<br />
AGROMYZIDAE<br />
Melanagromyza<br />
eupatoriella<br />
CEClDOMYllDAE<br />
Clinodiplosis sp.<br />
Perasphondylia<br />
reticulata<br />
TEPHRlTlDAE<br />
Procecidochares sp.<br />
Lepidoptera<br />
BUCCULATRICIDAE<br />
Bucculatrix sp.<br />
NYMPHALIDAE<br />
Actinote anteas<br />
PYRALIDAE<br />
Mescinia parvula<br />
stem<br />
stem<br />
leaf miner<br />
stem<br />
shoot borer<br />
shoot galls<br />
bud galls<br />
stem galls<br />
leaf miner<br />
leaf<br />
shoot borer<br />
moderate<br />
minor<br />
moderate<br />
great<br />
great<br />
great<br />
great<br />
moderate<br />
minor<br />
great<br />
great<br />
Trinidad<br />
all Americas<br />
prefers shade Trinidad<br />
Trinidad<br />
cage mating West Indies, S. America<br />
rearing Trinidad<br />
cage rearing all Americas<br />
parasites Americas<br />
Mexico<br />
cage mating Trinidad, Costa Rica<br />
cage mating all Americas<br />
causes conspicuous leaf lesions. Pseudocercospora eupatorii-formosani is reported to be<br />
common and damaging on C. odorata in Brunei, but is widespread already in South and<br />
Southeast Asia (Chacko 1988, Evans 1987, Peregrine and Ahmad 1982).<br />
Attempts at biological control<br />
Four insects have been released for biological control, the weevil Apion brunneoni-<br />
grurn, the fly Melanagromyza eupatoriella and two moths Mescinia parvula and<br />
Pareuchaetes pseudoinsulata (Table 4.4.4). Of these, only the last has become estab-<br />
lished, fairly readily in Sri Lanka, Guam and other Micronesian islands, but with some<br />
difficulty in India and Sabah (Malaysia) and it 'has since spread unaided to the<br />
Philippines and Brunei. It failed to become established in Thailand, Ghana, Nigeria and<br />
South Africa. It has produced spectacular defoliation and death of many plants in Guam
40 Biological Control of Weeds: Southeast Asian Prospects<br />
Table 4.4.2 Additional potential biological control agents for C. odorata: arthropods.<br />
Species Part attacked Country found<br />
Coleoptera<br />
CERAMBYCIDAE<br />
Aerenica hirticornis stem borer Trinidad, Bolivia,<br />
Brazil, Argentina<br />
CURCULIONIDAE<br />
Baris sp.<br />
Centrinaspis sp.<br />
Diptera<br />
CEClDOMYllDAE<br />
Asphondylia corbulae<br />
Clinodiplosis eupatorii<br />
Clinodiplosis sp.<br />
Contarinia sp.<br />
Neolasioptera brickelliae<br />
Neolasioptera cruttwellae<br />
Neolasioptera eupatorii<br />
Neolasioptera frugivora<br />
TEPHRITIDAE<br />
Cecidochares jluminensis<br />
Procecidochares connexa<br />
flowers, leaf buds<br />
flowers, leaf buds<br />
flower galls<br />
leaf galls<br />
bud galls<br />
flowers (achenes)<br />
flowers (achenes)<br />
stem galls<br />
stem galls<br />
flowers (achenes)<br />
flowers<br />
stem galls<br />
Trinidad<br />
Trinidad, Costa Rica<br />
El Salvador, Trinidad<br />
Central America, Brazil, West<br />
Indies<br />
Trinidad<br />
Trinidad<br />
Trinidad<br />
Trinidad<br />
USA, Trinidad, Bolivia<br />
Trinidad<br />
Trinidad, SE Brazil<br />
Mexico, Brazil, Bolivia<br />
Lepidoptera<br />
ARCTllDAE<br />
Pareuchaetes aurata aurata leaves. buds Paraguay, Argentina,<br />
Pareuchaetes aurata aurantior<br />
Pareuchaetes arravaca<br />
Pareuchaetes insulata<br />
Pareuchaetes misantlensis<br />
Pareuchaetes sp.<br />
GELECHllDAE<br />
Dichomeris (= Trichotaphe) sp.<br />
leaves, buds<br />
leaves, buds<br />
leaves, buds<br />
leaves, buds<br />
leaves, buds<br />
leaf roller (see<br />
Cruttwell 1973b) Trinidad<br />
SE Brazil, Bolivia<br />
Amazon River<br />
Surinam and French Guiana<br />
Southern USA, Mexico,<br />
Caribbean, Colombia<br />
Mexico<br />
Mexico<br />
LYCAENIDAE<br />
Calephelis laverna leaves Brazil, Trinidad, Venezuela,<br />
Central America<br />
Acarina<br />
ERlOPHYlDAE<br />
Calacarus sp. shoots<br />
India
Table 4.4.3 Natural enemies of C. odorata: fungi.<br />
4.4 Chromolaena odorata 41<br />
Species Country found References<br />
Anhellia niger<br />
Cercospora sp.<br />
Cercospora eupatorii<br />
Cercospora eupatoriicola<br />
Cercospora eupatorii-odoratii<br />
Cionothrix praelonga<br />
Fusarium oxysporum f. sp. elaeidis<br />
Guignardia eupatorii<br />
Mycovellosiella perfoliata<br />
Phoma sp.<br />
Phomopsis eupatoriicola<br />
Phyllosticta eupatoriicola<br />
Pseudocercospora eupatorii-formosani<br />
Septoria sp.<br />
Septoria ekmaniana<br />
Trinidad, Tobago<br />
Peninsular Malaysia,<br />
Sabah<br />
North America, Cuba,<br />
Nepal, India, Ivory Coast<br />
India, Bangladesh<br />
Malaysia<br />
Dominica, Tobago,<br />
Venezuela<br />
Trinidad, Tobago<br />
Sri Lanka<br />
Trinidad, Tobago<br />
Trinidad, Tobago<br />
not recorded<br />
not recorded<br />
India, Myanmar, Thailand,<br />
Malaysia, Borneo, Brunei,<br />
Guam<br />
Trinidad, Tobago<br />
Ooi et al. 1991<br />
Singh 1980<br />
Chacko 1988, Evans 1987<br />
Chacko 1988, Evans 1987<br />
Chacko 1988, Evans 1987<br />
Chacko 1988, Evans 1987<br />
Oritsejafor 1986<br />
Chacko 1988, Evans 1987<br />
Ooi et al. 1991<br />
Ooi et al. 1991<br />
Chacko 1988, Evans 1987<br />
Chacko 1988, Evans 1987<br />
Chacko 1988, Evans 1987<br />
Russo 1985<br />
Ooi et al. 1991<br />
and striking but sporadic defoliation in Sri Lanka. In India, populations have built up but<br />
damage has seldom been great. Where established, it is heavily attacked by a range of<br />
predators and these are believed to have prevented successful establishment in several<br />
countries.<br />
The eriophyid mite Acalitus adoratus causes abnormal growth of the epidermal<br />
hairs on young leaves and stems of C. odorata. Although it was never purposely intro-<br />
duced, it was observed in Thailand in 1984 and the Philippines in 1987, but had probably<br />
been present for some years. It is also widespread in Java and Sumatra, but there is no<br />
information from other Indonesian islands. It is present in Yap and Palau in the Caroline<br />
Islands and was observed on Guam in November 1993 (R. Muniappan pers. comm.). It is<br />
not present in India and it is not known whether it is present in Sri Lanka (Cruttwell<br />
1977b, McFadyen 1991 b, 1993, Muniappan et al. 1988a).<br />
Further details follow of the situation in individual countries and of the biology of<br />
some of the more promising natural enemies.<br />
Asia<br />
INDIA<br />
In one study, 11 insects and 3 mites were found attacking C. odorata. All except<br />
Calacarus sp. (Eriophyidae) are polyphagous (Muniappan and Viraktamath 1986,<br />
Viraktamath and Muniappan 1992). Most eriophyid mites have a highly restricted host<br />
range, so it is not clear whether it may even have accompanied the weed from the
42 Biological Control of Weeds: Southeast Asian Prospects<br />
4.4.4 Introductions for the biological control of Chromolaena odorata.<br />
Species Country Liberated Result References<br />
Coleoptera<br />
BRENTHIDAE<br />
Apion brunneonigrum Ghana 1975 - Cock 1984a, 1985,<br />
Guam 1984 - Nafus & Schreiner 1989<br />
India 1972-83 - Chacko & Narasimham<br />
1988, Cock 1984, 1985<br />
Malaysia 1970 - Ooi et al. 1988a,b<br />
(Sabah)<br />
Nigeria 197CL75 - Cock 1984a, 1985<br />
Sri Lanka 197676 - Cock 1984a, 1985<br />
Lepidoptera<br />
ARCTllDAE<br />
Pareuchaetes aurata<br />
aurata<br />
Pareuchaetes<br />
pseudoinsulata<br />
South Africa<br />
Pohnpei<br />
Ghana<br />
Guam<br />
India<br />
Indonesia<br />
Kosrae<br />
Malaysia<br />
(Sabah)<br />
Northern<br />
Marianas<br />
Nigeria<br />
South Africa<br />
Sri Lanka<br />
Thailand<br />
Yap<br />
Julien 1992,<br />
Kluge & Caldwell 1993<br />
Esguerra et al. 199 1,<br />
Esguerra et al. 1994<br />
Muniappan et al. 1988b<br />
Cock 1985,<br />
Cock & Holloway 1982<br />
Julien 1992<br />
Nafus & Schreiner 1989,<br />
Seibert 1989<br />
Chacko & Narasimham<br />
1988, Cock & Holloway<br />
1982<br />
Chacko & Narasimham<br />
1988, Julien 1992,<br />
Muniappan et al. 1989,<br />
Satheesan et al. 1987<br />
McFadyen pers. comm.<br />
Esguerra et al. 1994<br />
Cock & Holloway 1982,<br />
Ooi et al. 1988a,b,<br />
Syed 1979a<br />
Nafus & Schreiner 1989,<br />
Seibert 1989<br />
Cock & Holloway 1982<br />
Julien 1992, Kluge 1991,<br />
Kluge & Caldwell 199 1<br />
Dharmadhikari et al. 1977<br />
Napompeth et al. 1988<br />
Marutani & Muniappan<br />
199 1 a, Muniappan et al.<br />
1988b<br />
PY RALl DAE<br />
Mescinia parvula Guam Nafus & Schreiner 1989
4.4 Chromolaena odorata 43<br />
Americas. In another study 21 polyphagous insects were recorded from C. odorata, of<br />
which the most widespread and numerous were Aphis fabae and A. spiraecola (Lyla and<br />
Joy 1992, Lyla et al. 1987). Some of these same species are included amongst the 31<br />
insects and 9 mites recorded on Chromolaena by Chacko and Narasimham (1988).<br />
Pareuchaetes pseudoinsulata from Trinidad was cleared of a nuclear polyhedral<br />
virus and mass reared. It was first released in 1973 at several sites in Karnataka, but no<br />
establishment occurred. Observations suggested that two ants, Myrmicaria brunnea and<br />
Oecophylla smaragdina, were major predators (Cock and Holloway 1972). P. pseudoin-<br />
sulata from Sri Lanka (where it had been sent and had already become established) was<br />
next released and appeared to be doing well until unexpectedly wiped out by virus (Cock<br />
1985). However, further material from Sri Lanka was laboratory reared and 36000 larvae<br />
and 1000 adults released from 1984 onwards, this time in Kerala. This procedure resulted<br />
in field establishment (Chacko and Narasimham 1988, Joy et al. 1993, Muniappan et al.<br />
1989, Satheesan et al. 1987). Most recently, the establishment of P. pseudoinsulata at<br />
Sullia Taluk in Karnataka State and defoliation of Chromolaena thickets over about 1000<br />
km2 was reported in December 1992 (R. Muniappan pers. comm. 1993). However the<br />
overall performance of the moth has been unsatisfactory (Joy et al. 1993).<br />
Small releases of the weevil Apion brunneonigrum have been made since 1972, but<br />
establishment has not resulted (Cock 1985, Ooi et al. 1991).<br />
SRI LANKA<br />
R pseudoinsulata was received from India in 1973 and about 2000 larvae released in a<br />
coconut estate in the North Western Province. Six months after release spectacular defo-<br />
liation was observed of a hectare of previously impenetrable growth of C. odorata. In<br />
addition to leaves, terminal buds and tender stems were being consumed. Further releas-<br />
es were made and two years later it was estimated that some 800 ha of C. odorata had<br />
been defoliated (Dharmadhikari et al. 1977). Since then sporadic, heavy defoliation has<br />
mainly occurred at the beginning of the dry season at the time of flowering. This has<br />
caused great damage and, at times, death of the weed. However R pseudoinsulata popu-<br />
lations fluctuate considerably, due in no small measure to natural enemies. Young larvae<br />
are taken by birds and predatory Sycanus bugs. They are also parasitised by the braconid<br />
Apanteles creatonoti and the tachinid Exorista sp. (Kanagaratnam 1976). In one series<br />
of experiments from 63 to 100% of pupae were consumed by ants, termites and lizards<br />
(Mahindapala et al. (1980). Perera (1981) fed P. pseudoinsulata larvae on C. odorata<br />
leaves dipped in 32P labelled sodium orthophosphate, transferred them to C. odorata in<br />
the field and collected predators from pitfall traps. Several carabids and a histerid<br />
showed no radioactivity, nor did the ants Odontomachus simillimus and Diacamma<br />
rugosum which were observed carrying away treated larvae to their nests, indicating that<br />
they do not feed on the larvae soon after capture. There was no unusual preponderance<br />
of predatory wasps, but birds were observed picking up larvae so it is likely that they<br />
were the cause of the sudden decline in larval population (P.A.C.R. Perera pers. comm.<br />
1993).
44 Biological Control of Weeds: Southeast Asian Prospects<br />
How effective the moth is as a control agent is yet to be determined. However,<br />
Perera (1981) calculated that a t? pseudoinsulata larva from hatching to pupation con-<br />
sumes an average of 184.6 cm2 of leaf. Based on measurements of a heavy growth of C.<br />
odorata, there are 22.42x lo8 cm2 of leaf area per ha requiring about 12 million larvae to<br />
produce defoliation. Assuming an average egg production of 200 per female moth and a<br />
1 : 1 sex ratio, 12 million larvae could be produced in two generations (3 months) with a<br />
release of 600 to 700 females. Cock and Holloway (1982) have suggested that there is a<br />
better climate match between Sri Lanka and Trinidad than for most of the other countries<br />
where the moth has been released.<br />
Apion brunneonigrum were released between 1974 and 1976 and, two months after<br />
release, were seen on flower heads but have not been recovered since (Cock 1985,<br />
Kanagaratnam 1976, Ooi et al. 1991).<br />
Southeast Asia<br />
BRUNEI<br />
Although no releases of P. pseudoinsulata have been made, two females were trapped in<br />
the early 1980s, presumably having resulted from the colonies established in neighbour-<br />
ing Sabah (Malaysia) (Cock 1985).<br />
INDONESIA<br />
An aphid has been observed to attack young shoots and cause leaf curl of C. odorata.<br />
Work on biological control of the weed was initiated in 1991 with the introduction of P.<br />
pseudoinsulata to Sumatra, but there is no information on the outcome. There is a current<br />
project (1993) under R.E. McFadyen to study the host specificity of the tephritid fly<br />
Procecidochares connexa and either the moth Mescinia parvula, the stem boring<br />
Melanagromyza eupatoriella or the butterfly Actinote anteas with a view, if judged safe<br />
to do so, to liberation in Indonesia and the Philippines (R.E. McFadyen pers. comm.,<br />
Tjitrosoedirdjo 1991, Tjitrosoedirdjo et al. 199 1).<br />
MALAYSIA (SABAH)<br />
Aphis spiraecola attacks young shoots of C. odorata and causes leaf fall (Bennett and<br />
Rao 1968).<br />
t? pseudoinsulata was introduced from India to Sabah in 1970 and releases made<br />
between 1970 and 1974 of over 4000 eggs, 40 000 larvae and 700 adults. Temporary<br />
establishment occurred in two areas in 1973 and 1974, but both colonies then appeared to<br />
die out over the next couple of years. This was considered to be due to general predators,<br />
such as ants (Cock and Holloway 1982). However, in 1983 and 1987, pockets of larvae<br />
appeared scattered over Sabah and often distant from the sites of original release (Ooi et<br />
al. 1988a,b). There is a good climate match between Sabah and Trinidad, which may<br />
explain the establishment (Cock and Holloway 1982).<br />
Small releases of A. brunneonigrum were made in 1970 and recoveries were report-<br />
ed a year later, but there is no indication that the weevil has survived (Ooi et al. 1988a,b,<br />
Syed 1973, 1975, 1979a,b).
4.4 Chromolaena odorata 45<br />
PHILIPPINES<br />
Aphis gossypii, A. spiraecola (= A. citricola) and the tortricid Homona coflearia were<br />
found attacking C. odorata (Torres 1986) and the total of natural enemies increased to 11<br />
by 8 additional (unspecified) insects (Torres 1988). Although it had not been intentional-<br />
ly introduced, numerous larvae of P. pseudoinsulata were discovered in 1985 in a limited<br />
area near the coast of Palawan. They were feeding on the leaves and stems of C. odorata<br />
under coconut trees and along roads, but surveys elsewhere at the time revealed no evi-<br />
dence of P. pseudoinsulata larvae (Aterrado 1986a,b, Torres and Paller 1989). However<br />
P. pseudoinsulata was discovered later in Zamboanga City, Bohol and northern Leyte<br />
provinces in the Visayas islands (Aterrado and Talatala-Sanico 1988).<br />
The eriophyid mite Acalitus adoratus was discovered in the Philippines in 1987<br />
(McFadyen 1 99 1 b).<br />
THAILAND<br />
A number of insects were found attacking C. odorata: the aphids Aphis craccivora,<br />
A. gossypii and A. spiraecola, the weevil Hypolixus trunculatus, a stem boring cicindelid<br />
larva and the arctiid moth Amsacta lactinea. They were causing little damage<br />
(Napompeth 1990a,b, Napompeth et al. 1988, Napompeth and Winotai 199 1).<br />
P. pseudoinsulata was introduced from Guam from 1986 to 1988 but, despite<br />
repeated field releases in 1987 and 1988, did not become established. The shoot miner<br />
Melanagromyza eupatoriella was introduced from Trinidad in 1978, but could not be<br />
reared and was not released.<br />
The mite Acalitus adoratus, detected in 1984, has since spread to all C. odorata<br />
infested areas, but is not having a significant effect on the weed.<br />
VIETNAM<br />
Infestations of Aphis craccivora and A. gossypii have been recorded on C. odorata, but<br />
no releases of biological control agents have been made (Napompeth and Hai 1988).<br />
Pacific<br />
GUAM<br />
P. pseudoinsulata was introduced from India and Trinidad, mass reared and first released<br />
in Guam in 1984 and later in the Northern Marianas. Initially late instar larvae were<br />
released in batches of up to 800, but were heavily attacked by ants, spiders, toads and<br />
other general predators and failed to become established. Next, groups of 500 or more<br />
adult moths were released at a number of sites, resulting in establishment in all release<br />
areas. Populations built up rapidly, defoliation of Chromolaena soon followed and almost<br />
all plants were stripped. Shoots arising from the crowns were also attacked as they<br />
appeared and, within a year, over 90% of the plants were killed. The moth spread rapidly<br />
and by 1987 had reached almost all infested areas of Guam. Eventually more than<br />
25 000 ha of the weed had been defoliated (Muniappan 1988c, Nafus and Schreiner 1989,<br />
Seibert 1989). A parasitoid Exorista xanthaspa (= E. civiloides) caused up to 30% mor-<br />
tality and predation by ants, spiders, toads and lizards occurred (Seibert 1989).
46 Biological Control of Weeds: Southeast Asian Prospects<br />
It was observed that the feeding of P. pseudoinsulata larvae caused the leaves of C.<br />
odorata to turn yellow, an effect that could not be produced by simply applying larval<br />
excreta to the plant. Yellow leaves were tougher and had a higher level of nitrate and,<br />
when larvae were forced, much against their preference, to feed on yellow leaves, they<br />
exhibited slow growth and high mortality. Furthermore, larvae continued to feed on yel-<br />
low plants both by day and night (exposing them to daytime predators), whereas on green<br />
plants they fed at night and hid at ground level by day (Marutani and Muniappan 1991 b).<br />
Interestingly, the yellow plants appear to lose their allelopathic properties and hence this<br />
major aid to dominance over other vegetation. The yellowing is reversible if the insects<br />
are removed (McConnell et al. 1992, Muniappan and Marutani 1992).<br />
Three additional insects were released to aid in the control of Chromolaena, but<br />
there is no evidence of establishment. Apion brunneonigrum was released early in 1984<br />
at the beginning of the dry season when the above ground growth of Chromolaena dies<br />
back. Because of the unsuitable condition of the host plants the beetle was not expected<br />
to become established. Small numbers of Mescinia parvula were released late in 1984<br />
and again late in 1986 (Seibert 1989). The mite Acalitus adoratus appeared in Guam in<br />
1993 (R. Muniappan pers. comm.).<br />
Larvae of the pyralid moth Eucampyla etheiella were observed attacking young<br />
flower buds and mature flowers and causing extensive damage. Larvae were parasitised<br />
by the eulophid Elachertus sp. and the elasmid Elasmus sp. (Marutani and Muniappan<br />
1 990).<br />
NORTHERN MARIANAS (ROTA, TINIAN, SAIPAN, AGUIJAN)<br />
P. pseudoinsulata has been established from liberations in 1986 and 1987 on all of these<br />
islands (Muniappan et al. 1989, Nafus and Schreiner 1989).<br />
Federated States of Micronesia<br />
KOSRAE<br />
Monthly releases of P. pseudoinsulata larvae in batches of 1000 to 4000 were made from<br />
early 1992 in sunny areas and defoliation of C. odorata was observed six months later.<br />
Predators were less active in sunny than in shady locations (Esguerra et al. 1994).<br />
PA LA U<br />
Although no releases of biological control agents have been made, the mite Acalitus ado-<br />
ratus was found to be present (Muniappan et al. 1988b).<br />
POHNPEI<br />
P. pseudoinsulata larvae were introduced from Guam in 1988, some liberated and others<br />
mass reared during which both larvae and adults were released until 1992. In four release<br />
sites extensive feeding injury and heavy defoliation of C. odorata was observed in 1991<br />
and populations persisted in 1992 in burnt areas where Siam weed was regenerating from<br />
root stocks (Esguerra et al. 1994). Heavy predation, especially in shaded conditions, was<br />
observed on all stages by ants, spiders, birds and lizards (Esguerra et al. 1991, 1994).
4.4 Chromolaena odorata 47<br />
YAP<br />
C. odorata was first reported in 1987. P. pseudoinsulata was released in 1988 at 14 dif-<br />
ferent sites, but failed to become established except at one location where only 100 lar-<br />
vae and 104 adults had been released (Muniappan et al. 1988b). It eventually disappeared<br />
at this site. However releases of 500 larvae in September and October and several hun-<br />
dred in December 1990 to June 1991 resulted in establishment (Marutani and Muniappan<br />
1991a). As on Guam and Rota, larvae of Eucampyla etheiella were found causing exten-<br />
sive damage to buds and mature flowers (Marutani and Muniappan 1990). The eriophyid<br />
mite Acalitus adoratus was found attacking C. odorata late in 1988, although it was not<br />
observed during a survey of the weed in May of that year (Muniappan et al. 1988b).<br />
Africa<br />
GHANA<br />
P. pseudoinsulata from India was used to establish a culture and releases were made<br />
between 1973 and 1978 in a variety of habitats including oil palm plantations. Although<br />
small amounts of leaf damage were observed shortly after release, no recoveries were<br />
made. Failure to establish was ascribed to predators, in particular to ants (Cock and<br />
Holloway 1982).<br />
One small release of Apion brunneonigrum was made in 1975, but it failed to<br />
become established (Cock 1985).<br />
NIGERIA<br />
P. pseudoinsulata shipped from Ghana between 1973 and 1978 were released, but no<br />
establishment occurred (Cock and Holloway 1982).<br />
A. brunneonigrum was sent from Trinidad from 1970 to 1975, but there is no record<br />
of establishment (Cock 1985).<br />
SOUTH AFRICA<br />
Disease-free adults of P. pseudoinsulata originating from Guam were released in batches<br />
of 500 to 1000 at 10 sites in Natal in 1989, but there are no signs of establishment. Very<br />
heavy egg predation (up to 82%) by ants and chrysopids was observed (Kluge 1991,<br />
Kluge and Caldwell 1991). P. pseudoinsulata has been obtained from Florida where the<br />
climate is similar to that in Natal and where there is a rich ant fauna. It is (as of 1991) to<br />
be released as soon as laboratory cultures of larvae have been cleared of microsporida.<br />
The larvae of another arctiid moth, Pareuchaetes aurata aurata, from Chromolaena<br />
jujuensis in Argentina were found to feed voraciously and complete their development on<br />
C. odorata. Females scatter their eggs around the base of the host plant and it is hoped<br />
that this will help to overcome the problem of ant predation. After specificity testing it<br />
has been released in Natal, but no further information is available (Kluge and Caldwell<br />
1993).<br />
'A laboratory culture of the butterfly Actinote anteas has been established with<br />
material collected in Costa Rica and host testing is to commence (Kluge and Caldwell<br />
1991).<br />
Work is also in progress on the host specificity of the weevil Rhodobaenus
48 Biological Control of Weeds: Southeast Asian Prospects<br />
cariniventris and a leaf spot disease caused by Septoria sp. (Kluge and Morris 1992).<br />
Major natural enemies<br />
Acalitus adoratus Acarina: Eriophyidae<br />
Recorded originally from Trinidad, Florida, Brazil and Bolivia, it appeared without spe-<br />
cial assistance in Thailand, Philippines, Indonesia (Java and Sumatra), Caroline Is (Yap,<br />
Palau) Guam and southern China. It was not present in India in the mid 1980s nor in Sri<br />
Lanka or West Africa (McFadyen 1993).<br />
These tiny mites (0.1 4 to 0.1 8 mm long) usually live on the lower surface of leaves.<br />
Their feeding induces abnormal growth of the epidermal hairs, resulting in the formation<br />
of erineum patches, the term given to areas covered with dense twisted hairs amongst<br />
which the mites live. These appear as white patches on the leaves, usually 0.5 to 3 mm in<br />
diameter, and the whole leaf surface may be affected. The patches often turn yellow on<br />
older leaves. The nymphs and adults feed, and the eggs are laid, between the epidermal<br />
hairs. Particularly heavy infestations develop in dry and exposed situations and, although<br />
the damage is not spectacular, heavy attack stunts, distorts and slows growth, thereby<br />
reducing competitiveness. Tests indicated that, as with many other eriophyid mites,<br />
A. adoratus is host specific.<br />
When infested leaves senesce, A. adoratus leave the erineum patches and are preyed<br />
upon by other mites and by the larvae of a cecidomyiid fly, Arthrocnodax meridionalis<br />
(Cruttwell 1977b, McFadyen 199 1 b, Muniappan et al. 1988a,b).<br />
It is suggested that A. adoratus was accidentally introduced to Sabah when field-<br />
collected adults of the weevil Apion brunneonigrum in Trinidad were released directly in<br />
the 1970s; and that it has since spread naturally and on leaves of C. odorata used as<br />
packing material around fruit and other produce (McFadyen 1993).<br />
Actinote anteas Lepidoptera: Nymphalidae<br />
Recorded from Costa Rica and Trinidad. The host specificity of this acraeinine defoliator<br />
is being examined in South Africa (Kluge and Caldwell 1991).<br />
Apion brunneonigrum Coleoptera: Apionidae<br />
This weevil has been recorded from Trinidad, Venezuela and Argentina. Small releases<br />
were made in West Africa, India, Sri Lanka, Sabah and the Marianas, but the weevil per-<br />
sisted only in Sabah and then apparently only briefly. The reasons for these failures have<br />
not been investigated.<br />
Cruttwell (1973a) studied its biology and host specificity and found that it would<br />
feed and develop only on C. odorata and C. ivaefolia. It has never been recorded damag-<br />
ing economic plants either in Trinidad or South America. The life history is closely linked<br />
with the development of its host, the adults becoming reproductively mature at the time<br />
that the plant produces young flower buds which provide food for egg maturation. Eggs<br />
are deposited in the developing flower heads and larvae feed within the flower heads,<br />
destroying the seeds. Pupation occurs in the flowerheads and, until the next flowering,<br />
adults feed on tender growth, usually in shaded situations, and may do considerable dam-
4.4 Chromolaena odorata 49<br />
age (Cock 1984a). An individual larva destroys 30 to 60 seeds during development and<br />
the ovipositing female many young flowers. A. brunneonigrum thus has potential for<br />
causing considerable damage, particularly in lightly shaded conditions.<br />
Aulacochlamys sp. Coleoptera: Chrysomelidae<br />
Widespread and occasionally abundant in Trinidad, where it causes moderate damage.<br />
Eggs are laid singly in a cylindrical ribbed case formed from faeces. These cases form<br />
the apex of a conical larval case which is enlarged as the larva grows. Larvae feed on the<br />
surface of stems and leaf petioles. Mature larvae attach the 3.5 to 3.7 mm long case to a<br />
stem and pupate inside. Adults emerge one to two weeks later. The small black adults<br />
(1.8 to 2.5 mm long) feed on the surface of stems and petioles. No parasitoids are known<br />
(McFadyen 1988a).<br />
Bucculatrix sp. Lepidoptera: Bucculatricidae<br />
Recorded from C. odorata in Mexico and Chromolaena jujuensis (= Eupatorium hooke-<br />
rianum) in Argentina. Larvae are solitary leaf miners and pupate in the mines (McFadyen<br />
1988a).<br />
Chlamisus insularis Coleoptera: Chrysomelidae<br />
Recorded from Mexico, Panama and Trinidad. The life history is similar to that of<br />
Aulacochlamys sp., but this species is somewhat larger. The mature larval case is conical<br />
with a rough surface and 6 to 7 mm long. The adults are black with golden markings and<br />
3.3 to 4.3 mm long. Adults are known to feed on C. odorata, C. ivaefolia and Bidens<br />
pilosa. A black, solitary eulophid egg parasitoid is known (McFadyen 1988a).<br />
Clinodiplosis sp. Diptera: Cecidomyiidae<br />
Recorded from C. odorata and C. ivaefolia in Trinidad. Up to three larvae at a time live<br />
between the bud leaves of stem tips or axillary buds, destroying tissue and preventing<br />
growth. A gall is formed by the slight swelling of the bud leaves which become red and<br />
densely covered with hairs. Mature larvae drop to the ground and pupate just below the<br />
soil surface. Adults emerge 11 to 18 days later. This gall midge is abundant and wide-<br />
spread in Trinidad, breeds throughout the year and causes considerable damage to C.<br />
odorata (McFadyen 1988a).<br />
Mescinia parvula Lepidoptera: Pyralidae<br />
Recorded from Trinidad; similar larvae were found on C. odorata in Mexico and Brazil<br />
and on C. jujuensis in Argentina. A few individuals were released on Guam in 1984, but<br />
there has been no evidence of establishment.<br />
Ovipositing females select leaves with dense hairs (in effect young leaves) with the<br />
result that developing buds are nearby. Eggs are laid individually amongst the epidermal<br />
hairs on the underside of the young leaves and hatch in 5 to 6 days. Young larvae move<br />
to a terminal or axillary bud and several may enter the same bud. They bore down the<br />
stem destroying meristematic tissue and preventing growth. Larvae may leave a stem and<br />
enter a new bud. After 13 or so days larvae leave the stem to spin a flimsy cocoon, either<br />
attached to the plant or among ground litter, in which they pupate. Adults emerge 10 to
50 Biological Control of Weeds: Southeast Asian Prospects<br />
11 days later and live up to 6 days. Attempts to induce mating in cages in Trinidad were<br />
unsuccessful. In specificity tests larvae fed on only a few Asteraceae other than C. odora-<br />
ta and C. ivaefolia but, with the exception of 1 out of 30 larvae placed on Dahlia, no<br />
development was ever completed. Over a three year period in Trinidad, no oviposition or<br />
attack was observed on Dahlia plants growing near C. odorata which was frequently<br />
attacked by M. parvula. Furthermore, since Dahlia leaves are not hairy, it is most unlike-<br />
ly that M. parvula would ever oviposit on them. Larvae in Trinidad are attacked by eight<br />
hymenopterous and one tachinid parasitoid and, if freed from these, might do consider-<br />
ably more damage to C. odorata (Cruttwell 1977a).<br />
Pareuchaetes aurata aurata Lepidoptera: Arctiidae<br />
This subspecies occurs in south-eastem Brazil, Paraguay and northern Argentina at lati-<br />
tudes (26" to 30°S) similar to those of Natal, South Africa. It has an average life cycle of<br />
30 days at 26' to 29OC and 58 days at 22O to 25OC. Its larvae are nocturnal feeders and<br />
shelter at the base of plants during the day. In the laboratory, pupation occurred in a flimsy<br />
cocoon spun between leaves on the plant. Mating may occur on the night of emergence<br />
and an average of 242 eggs are laid over the next eight days. These are laid singly on the<br />
ground and, in the laboratory, newly-hatched first instar larvae were able to walk up about<br />
2m of stem to commence feeding. In the field ?! aurata aurata is found in shaded habitats<br />
near surface water.<br />
In Argentina, larvae and pupae are infected with a microsporidan disease (Nosema<br />
sp.), up to 20% of larvae are parasitised by a complex of braconid, chalcidid and tachinid<br />
parasitoids and all stages are subject to attack by predatory ants.<br />
The usual host plant of ?! aurata aurata is Chromolaena jujuensis, but it has been<br />
successfully reared for more than 10 generations on C. odorata. In the field it has never<br />
been recorded as a pest on any of the many commercially important crops grown in its<br />
natural area of distribution. It was liberated in Natal (South Africa) in 1990 (Kluge and<br />
Caldwell 1993).<br />
Pareuchaetes pseudoinsuhta Lepidoptera: Arctiidae<br />
This moth is native to Trinidad, Tobago and the north-eastern coast of Venezuela. It has<br />
become established in Brunei, Guam, India, Philippines, Sabah, Sri Lanka, the Northern<br />
Marianas and Yap, but has failed to do so in Ghana, Nigeria, South Africa and Thailand. It<br />
was previously misidentified first as Ammalo arravaca and then as A. insulata, which is a<br />
closely related but distinct species (Cock and Holloway 1982).<br />
The moth, which lives up to about 10 days, lays 150 to 250 eggs (maximum 580) in<br />
groups attached to the lower surface of the leaves of C. odorata. Larvae feed on the leaves<br />
and are gregarious until the 3rd instar, but then disperse. From the 4th instar on they feed<br />
at night, hiding by day amongst debris at the base of the plant, where they later pupate.<br />
The life cycle varies from 40 to 60 days and breeding occurs throughout the year. Host<br />
specificity studies in Trinidad showed that development occurred only on Chromolaena<br />
ivaefolium, C. microstemon and C. odoratum. In addition larvae developed, but only as far<br />
as the 3rd instar, on Ageratum conyzoides (Bennett and Cruttwell 1973), although in Sri<br />
Lanka, adults were produced on this weed. However their eggs had a somewhat lower
4.4 Chromolaena odorata 51<br />
hatchability than those from adults bred on C. odorata (Mahindapala et al. 1980). A high<br />
degree of host specificity has since been confirmed by others (e.g. Ahmad and Thakur<br />
1991, Sankaran and Sugathan 1974, Syed 1979a) and no damage to plants other than to C.<br />
odorata has been reported either in the Americas or in the overseas countries where it has<br />
become established.<br />
In Trinidad the eggs are parasitised by a scelionid wasp and the larvae by five<br />
species of tachinid fly. A nuclear polyhedrosis virus also affects the larvae (Bennett and<br />
Cruttwell 1973) and may have been responsible for breeding difficulties in some overseas<br />
countries, although other countries have experienced no problems in establishing cultures.<br />
Pentispa explanata Coleoptera: Chrysomelidae<br />
This hispine beetle is recorded on C. odoratum from Mexico to Colombia and from<br />
Venezuela on Pithecoctenium sp. (Bignoniaceae). In Trinidad adults are widespread on C.<br />
odorata and C. ivaefolia, but would not feed on Pithecoctenium echinatum.<br />
Eggs are inserted singly under the leaf epidermis and covered with a faecal plug.<br />
Larvae hatch after about 12 days and form irregular blotch mines which expand to 2 to 3<br />
cm in diameter 20 to 25 days later when larvae are mature. Pupation occurs in the mine<br />
and adults emerge 5 to 8 days later. Adults disperse and feed on the underside of the<br />
leaves producing characteristic scars. There is one generation a year. Larvae are para-<br />
sitised by a solitary ectoparasitic elasmid Austelasmus sp. and are taken by predatory<br />
wasps (McFadyen 1988a).<br />
Perasphondylia reticulata Diptera: Cecidomyiidae<br />
This gall fly is recorded from C. odorata and C. ivaefolia in Trinidad, Brazil and Bolivia<br />
and from C. odorata and Eupatorium sp. in El Salvador.<br />
Larvae occur singly in a hollow pear-shaped gall, 7 to 9 mm long, in stem tips and<br />
axillary buds. P. reticulata causes considerable damage but is generally uncommon and<br />
confined to the cooler valleys in Trinidad. However it is commoner in Brazil and Bolivia.<br />
It is attacked by several parasitoids in Trinidad and Bolivia (McFadyen 1988a).<br />
Procecidochares connexa Diptera: Tephritidae<br />
This gall fly is recorded from Mexico, Brazil and Bolivia.<br />
Eggs are inserted in the tip of the stem and abnormal growth commences even<br />
before they hatch. One to seven larvae feed in separate curved tunnels in the developing<br />
gall. Mature larvae pupate in the tunnel behind an epidermal window through which the<br />
adult emerges later. The galls slow and distort growth and cause moderate damage.<br />
Larvae are parasitised by a number of wasps throughout their range (McFadyen<br />
1988a).<br />
Rhodobaenus cariniventtis Coleoptera, Curculionidae<br />
This weevil is recorded from Trinidad and USA. Eggs are deposited in a slit between two<br />
rows 'of punctures encircling the stem, which result in wilting of the stem tip. On hatch-<br />
ing, larvae feed for a few days in the wilted portion then tunnel into the sound tissue<br />
below the punctures. Bennett (1955) reports that, after a month, they pupate in the stem<br />
at the base of the plant and adults emerge 10 days later. However, McFadyen (1988a)
52 Biological Control of Weeds: Southeast Asian Prospects<br />
states that, when mature, the larva cuts off from the hollow tip of the stem a piece about<br />
2cm long containing it. This falls to the ground, where the open ends are plugged with<br />
frass, and pupation occurs. Adults feed on stems and petioles of Bidens pilosa,<br />
Chromolaena inulaefolium, C. ivaefolia and C. odorata (all Asteraceae). Larvae tunnel in<br />
the stems of all these except B. pilosa (McFadyen 1988a). Cruttwell (1974) suggested<br />
that the feeding of adults might be insufficiently restricted, but the situation merits fur-<br />
ther investigation.<br />
R. cariniventris is parasitised in Trinidad by an external egg parasitoid Euderus sp.<br />
(Eulophidae) (Bennett 1 955).<br />
Comments<br />
The genus Chromolaena belongs to the tribe Eupatorieae (Table 4.3.4), which is mainly<br />
of American origin. There are no crop plants or important ornamentals in this tribe.<br />
However, it does contain the major weeds Ageratum conyzoides (4.1) and Mikania<br />
micrantha (4.14) and several less important species: Ageratina altissima in eastern USA,<br />
A. adenophora and A. riparia in India to southern China, Australia, Hawaii and South<br />
Africa and Austroeupatorium inulaefolium in Indomalaysia and Sri Lanka. There are 129<br />
species of Chromolaena, all from Central and South America. Chromolaena ivaefolia<br />
and C. laevigata are widespread and occasionally weedy in the Americas, but only C.<br />
odorata has spread elsewhere (McFadyen 1991a). These relationships suggest that many<br />
of the insects that attack C. odorata and its close allies are likely to be sufficiently host<br />
specific to be considered for classical biological control.<br />
There has been discussion concerning the possible reasons for Pareuchaetes<br />
pseudoinsulata establishing fairly readily in Sri Lanka and Guam, with difficulty in India<br />
and Sabah and not at all in Africa (e.g. Cock 1984a, Cock and Holloway 1982, Seibert<br />
1989). The desirability is rightly emphasised of matching, where possible, the climate of<br />
the area from which it (or any other biological control agent) is collected with that of the<br />
area in which it is to be released. Since P. pseudoinsulata has no diapause and breeds all<br />
year round, it will at least experience great difficulty in bridging (or find it impossible to<br />
bridge) the gap created by almost complete leaf loss of C. odorata in areas where there is<br />
a severe and long dry season. However, if this were the only problem, the moth should be<br />
able to establish itself at least briefly before being eliminated by starvation: this sequence<br />
has not, however, been documented. What, however, has been widely reported is the very<br />
high level of predation on eggs, larvae and pupae, in particular by ants and spiders, but<br />
also by other invertebrate and vertebrate predators (e.g. Kluge 1991, Kluge and Caldwell<br />
1991). It seems probable that massive predation has been the cause of rapid demise of<br />
many releases. Thus, release sites should be chosen (or treated) so as to minimise preda-<br />
tion. Although significant predation was also observed on Guam, the lower diversity of<br />
predators (and other organisms) on islands may well have contributed to the compara-<br />
tively ready establishment of P. pseudoinsulata there and on other Pacific islands.<br />
Furthermore, the release of significant numbers (500 or more) of adults rather than of<br />
eggs or larvae may have assisted in avoiding rapid elimination of the released material.
4.4 Chromolaena odorata 53<br />
The general vigour of the released insects and the presence or absence of microsporida or<br />
viruses would also play a crucial role in successful establishment and it is probable that<br />
these factors have not always been adequately considered.<br />
It is possible that the high level of predation and of parasitisation of the biological<br />
control agents will, in many areas, so lessen the potential each has to cause damage to C.<br />
odorata that the combined effects of several will be required to bring about an adequate<br />
and sustained reduction in its weediness. Fortunately, many potentially suitable insects<br />
are available for study.
84 Biological Control of Weeds: Southeast Asian Prospects<br />
Eleusine indica<br />
(after Holm et a/. 1977)
Map 4.8.1 Eleusine indica<br />
4.8 Eleusine indica 85<br />
Eleusine indica<br />
Eleusine indica is of African origin and, except for finger millet, E. coracana, is not<br />
closely related to graminaceous crop plants. Finger millet is a staple crop in India and<br />
some parts of Africa, but relatively unimportant or not grown elsewhere. Little is known<br />
about the insect or other enemies of E. indica in Africa and, elsewhere, almost all records<br />
are of pests with a wide host range. Because it is a major weed (5th) in Southeast Asia<br />
and is only distantly related to crop plants a search for specific natural enemies in Africa<br />
must be regarded as an attractive proposition.
86 Biological Control of Weeds: Southeast Asian Prospects<br />
4.8 Eleusine indica (L.) Gaertn.<br />
Poaceae<br />
crowsfoot grass, goose grass; sin ngo let kya, sin ngo myet (Myanmar), yah<br />
teenka (Thailand), smao choeung tukke (Cambodia) co miin triiu (Vietnam),<br />
rumput sambou (Malaysia), rumput belul8ng (Indonesia), sabung sabungan<br />
(Philippines)<br />
Rating<br />
+++ Viet, Msia, Sing, Indo, Phil<br />
24 ++ Myan, Thai, Laos, Camb<br />
+ Brun<br />
Origin<br />
Africa (Phillips 1972), replacing an alternative view that it was India (Holm et al. 1977,<br />
Waterhouse 1993a).<br />
Distribution<br />
Throughout the tropics, sub-tropics and temperate regions of the world, including Africa,<br />
Asia, SE Asia, Australia, the Pacific and the Americas.<br />
Characteristics<br />
E. indica is a tufted, annual, C4 grass attaining a height of 0.6 m. Its flower spikes mostly<br />
have 2 to 7 spikelets, producing a characteristic windmill-like appearance.<br />
Importance<br />
The genus Eleusine, contains nine annual or perennial grasses all native to Africa except<br />
for the South American E. tristachya (Hilu and Johnson 1992, Phillips 1972). It belongs<br />
to the subfamily Chloridoideae, which is but distantly related to all except one of the<br />
principal grain crops. That exception is finger millet, or ragi, E. coracana (2n = 36),<br />
which is believed to have arisen from E. indica (2n = 18) (Hilu and de Wet 1976, Hilu<br />
and Johnson 1992, Hiremath and Salimath 1992) and is an important staple cereal in<br />
India and some regions of eastern Africa (Rachie and Peters 1977). However, it is worth<br />
noting that E. coracana is regarded as a minor weed in some Southeast Asian countries<br />
(Thailand, Vietnam) (Waterhouse 1993a) and that it is nowhere important in this region.<br />
The genera Eleusine and Dactyloctenium are closely related.<br />
E. indica is an important weed in more than 60 countries in at least 46 crops and, in<br />
these, has the status of a serious weed in 30 countries and 27 crops. It was evaluated as<br />
the fifth worst weed in the world (Holm et al. 1977) and also rated fifth in a recent survey<br />
in Southeast Asia (Waterhouse 1993a). It was rated 15th in 1992 in the oceanic Pacific<br />
(Waterhouse, unpublished). It grows well in sunny or somewhat shaded places, in marsh-<br />
lands,. wastelands, roadsides, along borders of irrigated fields and canals, in lawns and in<br />
pastures, and prospers and is particularly troublesome on arable land. It ranges from near
4.8 Eleusine indica 87<br />
the seashore to an altitude of at least 2000 m and is a major problem in almost all forms<br />
of agriculture between the tropics of Capricorn and Cancer.<br />
E. indica grows and flowers well in all seasons and a single plant may produce more<br />
than 50000 small seeds, which move readily by wind, in mud on the feet of animals and in<br />
the tread of machinery. The seeds are eaten by wild and domestic animals and are occasion-<br />
ally grown for grain in Africa and India, but Eleusine coracana, finger millet, with some-<br />
what larger seeds is far better for this purpose. Although sometimes claimed to be palatable<br />
to grazing animals, crowsfoot grass becomes fibrous too early in the season to be a satis-<br />
factory pasture grass. The seed heads may contain high levels of cyanogenic glycosides<br />
and are believed to be responsible for occasional cases of stock poisoning (Everist 1974).<br />
Natural enemies<br />
Natural enemies restricted to the genus Eleusine and its close relatives might well be con-<br />
sidered for biological control of E. indica except in India or other regions where finger<br />
millet is an important cereal.<br />
E. indica is reported in the literature to be attacked by more than 50 insects, nema-<br />
todes, fungi, bacteria and viruses, all except 6 in continents other than Africa (Tables<br />
4.8.1 to 4.8.4). Further, with few exceptions, all of these organisms are known to have<br />
wide host ranges and to attack important agricultural crops. Indeed, of those recorded,<br />
only one cecidomyiid gall fly and possibly one or two fungi could be considered further<br />
for classical biological control. Figliola et al. (1988) consider that, where they already<br />
occur, two fungi, Bipolaris setariae and Magnaporthe (=Pyricularia) grisea hold<br />
promise as bioherbicides for E. indica, although their host range is a little too wide for<br />
classical biological control.<br />
It is not surprising that the organisms attacking an economic crop, finger millet,<br />
E. coracana, have been investigated in greater detail than those of a weedy relative.<br />
Finger millet is believed to have been domesticated in the East African highlands by<br />
3000 BC or earlier and archaeological data suggests that it may have been introduced to<br />
India as early as 2000 BC (Hilu et al. 1979). Since E. coracana and E. indica are closely<br />
related, Wapshere (1 990b) argues, probably correctly, that most or all of the more specific<br />
organisms infesting finger millet are also likely to attack E. indica. It is very disappoint-<br />
ing, therefore, that almost all of the natural enemies of finger millet so far recorded (again<br />
mostly from outside Africa) have wide to very wide host ranges and are not potential bio-<br />
logical control agents. The very few species that may prove to have a limited host range<br />
are shown in table 4.8.5. Wapshere (1990b) has listed 40 insects that attack E. coracana<br />
and at the same time belong to groups known to have species restricted to a single grass<br />
genus (and there are also many other insects from groups with a wider host range that<br />
attack E. coracana). In addition to the undescribed Orseolia gall midge attacking E. indi-<br />
ca in India, only three insects (two cecidomyiid gall midges, one from Uganda and one<br />
from Nigeria and an aphid from India), a nematode (Heterodera delvii) from India and a<br />
smut fungus (Melanopsichiurn (= Ustilago) eleusinis) may, if shown to attack E. indica<br />
also, prove to be sufficiently host specific to be considered for classical biological control.<br />
It is relevant that cecidomyiid gall flies are believed to have a high degree of specificity to<br />
their host grass genera (Barnes 1946, K.M. Harris pers. comm. 1993, Wapshere 1990a).
88 Biological Control of Weeds: Southeast Asian Prospects<br />
Comment<br />
It has been pointed out above that the majority of records for natural enemies of both<br />
E. indica and E. coracana come from outside Africa and that almost all of these organ-<br />
isms have a wide host range. Indeed, this is to be expected if both Eleusine species are of<br />
African origin. Except for any specific enemies that may have accompanied them, it is<br />
inevitable that they will be attacked in new countries by non-specific natural enemies<br />
that, hitherto, were attacking other plants. Of course, the possibility exists that natural<br />
enemy species in the new country may have evolved a degree of specificity in the four or<br />
five thousand years that the Eleusine species have existed outside Africa.<br />
It is significant that there has not so far been any detailed search in Africa for natur-<br />
al enemies of E. indica to establish whether or not adequately specific species occur<br />
there. A two year (or longer) survey of E. indica in several regions of Africa would prob-<br />
ably be required, together with observations on whether the organisms found attacking<br />
E. indica also attack E. coracana, nearby grasses or other plants. The relevant regions for<br />
study in Africa and Madagascar are indicated in map 4.8.2 based on the distribution data<br />
of Phillips (1 972).<br />
If (i) the African cecidomyiid gall midges (Contarinia (= Stenodiplosis) spp.)<br />
(Tables 4.8.1, 4.8.5) do not already occur in Southeast Asia (they are not known in<br />
Australia), (ii) they prove to be adequately host specific and (iii) the Ugandan species<br />
attacks E. indica in addition to E. coracana, they would appear to be the most promising<br />
of known species for introduction elsewhere. The undescribed species from northern<br />
Nigeria (Table 4.8.1) was collected from E. indica at Zaria in July 1959 and July 1960<br />
(K.M. Harris, pers. comm. 1993). Larvae of the Indian Orseolia sp. nr. fluviatilis proba-<br />
bly induce galls on young shoots of E. indica, so would affect vegetative growth rather<br />
than having a direct impact on seed production. It is as yet known only from India.<br />
To sum up, for an attempt at classical biological control of a grass weed, E. indica<br />
would appear to be the one with most positive factors combined except that, so far, few<br />
adequately specific, natural enemies are known. However, almost nothing is known about<br />
the natural enemies in Africa, not only its centre of origin but also that of the genus<br />
Eleusine. It would, indeed, be most surprising if several natural enemies having a restrict-<br />
ed host range were not discovered during a thorough survey there.<br />
Table 4.8.1 Natural enemies of Eleusine indica: insects.<br />
Species Country Portion Comments: References<br />
attacked other hosts<br />
Hemiptera<br />
APHlDlDAE<br />
Chaetogeoica India<br />
graminiphaga<br />
Geoica lucijiuga India<br />
beans and a Raychaudhuri 1983<br />
number of<br />
grasses<br />
also on rice, Eleusine Raychaudhuri et al.<br />
coracana and many 1978<br />
weeds including<br />
Cynodon dactylon,<br />
Cyperus rotundus<br />
I (continued on next page)
4.8 Eleusine indica 89<br />
Species Country Portion Comments: References<br />
attacked other hosts<br />
Hysteroneura setariae Hawaii rice, maize, wheat<br />
Schizaphis (= Toxoptera)<br />
graminum<br />
Sitobion avenae<br />
(= Macrosiphum<br />
granarium)<br />
Sitobion (= Macrosiphum)<br />
miscanthi<br />
Sregophylla (= Anoecia)<br />
querci<br />
Tetraneura basui<br />
Tetraneura<br />
nigriabdominalis<br />
(= T. hirsuta)<br />
CERCOPIDAE<br />
Prosapia (= Monecphora)<br />
bicincta fraterna<br />
CICADELLIDAE<br />
Nephotettix malayanus<br />
Nephotettix<br />
nigromaculatus<br />
(= N. nigropictus)<br />
Nephotettix virescens<br />
Recilia dorsalis<br />
DELPHACIDAE<br />
Laodelphax striatellus<br />
sorghum, sugar cane<br />
rice, sorghum, maize<br />
and a very wide<br />
host range<br />
rice, wheat, a very<br />
wide host range<br />
India on a very wide range<br />
of crop plants and<br />
weeds<br />
maize and several<br />
weeds<br />
India<br />
on rice, Echinochloa<br />
colona, Paspalum<br />
conjugatum and<br />
other weeds<br />
India<br />
rice, maize, sugarcane<br />
Eleusine coracana<br />
and a very wide<br />
range of weeds<br />
Cuba also on Paspalum<br />
notatum, Brachiaria<br />
subquadripara,<br />
Andropogon annulatus,<br />
Cynodon dactylon<br />
Philippines<br />
Philippines<br />
Philippines<br />
Philippines<br />
rice, many weeds<br />
rice, many weeds<br />
rice, many weeds<br />
rice, many weeds<br />
Beardsley 1962<br />
Patch 1939<br />
Patch 1939<br />
Raychaudhuri 1983<br />
Patch 1939<br />
Raychaudhuri 1983<br />
Patch 1939,<br />
Raychaudhuri 1983<br />
Plana et al. 1986<br />
Khan et al. 1991<br />
Khan et al. 1991<br />
Khan et al. 1991<br />
Khan et al. 1991<br />
China wheat, barley, oats,<br />
sorghum etc<br />
Zhang et al. 1981<br />
Peregrinus maidis India transmitter of<br />
Cherian and Kylasam<br />
Eleusine mosaic 1937, Patch 1939, Rao<br />
virus (see table 4.8.4);<br />
very wide host range<br />
et al. 1965<br />
Sogatella furcijera China can complete<br />
development also on<br />
17 other species of<br />
crops and weeds<br />
including rice, barley,<br />
wheat, Echinochloa<br />
crus-galli<br />
Huang et al. 1985<br />
(continued on next page)
90 Biological Control of Weeds: Southeast Asian Prospects<br />
Table 4.8.1 (continued)<br />
Species Country Portion Comments: References<br />
attacked other hosts<br />
LYGAEIDAE<br />
Blissus leucopterus<br />
Thysanoptera<br />
PHLAEOTHRIPIDAE<br />
Haplothrips ganglbaueri<br />
Diptera<br />
AGROMYZIDAE<br />
Liriomyza marginalis<br />
Pseudonapomyza spicata<br />
CEClDOMYllDAE<br />
Orseolia sp. nrjluviatilis<br />
Stenodiplosis sp.<br />
Lepidoptera<br />
ARCTllDAE<br />
Cnaphalocrocis medinalis<br />
Cnaphalocrocis<br />
(= Marasmia) patnalis<br />
Creatonotos (= Amsacta)<br />
gangis<br />
NOCTUI DAE<br />
Spodoptera ffugiperda<br />
PYRALI DAE<br />
Ostrinia furnacalis<br />
USA lower damages sorghum and Ahmad et al. 1984,<br />
stem many grasses including Lynch et al. 1987<br />
Cynodon dactylon and<br />
Dactyloctenium<br />
aegyptium, but<br />
particularly damaging<br />
to E. indica<br />
India rice, wheat, sorghum Ananthakrishnan &<br />
Thangavelu 1976<br />
N&S Panicum miliaceum,<br />
America Digitaria, Paspalum<br />
(primary host),<br />
Euchlaena<br />
Australia leaf maize, sugarcane,<br />
grasses<br />
India stem undescribed gall midge<br />
resembling (but not)<br />
the rice stem gall midge<br />
Orseolia<br />
(= Pachydiplosis)<br />
oryzae; no host other<br />
than E. indica known<br />
Nigeria seed undescribed species<br />
heads<br />
Spencer 1990,<br />
Spencer & Steyskal<br />
1986<br />
Kleinschmidt 1970<br />
Barnes 1954a,b, 1956,<br />
Gagnt 1985,<br />
Hegdekatti 1927,<br />
Rachie and Peters,<br />
1977<br />
K.M. Harris pers.<br />
comm. 1993<br />
Philippines leaf rice, many weeds Abenes & Khan 1990<br />
folder<br />
Philippines leaf rice, many weeds Abenes & Khan 1990<br />
folder<br />
Philippines leaves rice, many weeds Catindig et al. 1993<br />
USA<br />
Guam<br />
wide range of crops Pencoe and Martin<br />
and weeds 1982<br />
wide range of crops Schreiner et al. 1990<br />
and weeds
Table 4.8.2 Natural enemies of Eleusine indica: nematodes.<br />
Species<br />
Ditylenchus destructor<br />
Hirschrnaniella spinicaudata<br />
Meloidogyne sp.<br />
Meloidogyne arenaria<br />
Meloidogyne grarninicola<br />
Meloidogyne incognita<br />
Meloidogyne javanica<br />
Pratylenchus pratensis<br />
Pratylenchus zeae<br />
Rotylenchulus reniformis<br />
4.8 Eleusine indica 91<br />
Country Comments References<br />
South Africa<br />
Cuba<br />
China<br />
Cuba,<br />
Philippines,<br />
USA<br />
India<br />
Cuba, USA<br />
Brazil<br />
Hawaii<br />
S. Africa,<br />
Cuba<br />
Hawaii<br />
groundnut, several weeds<br />
has other weed hosts<br />
including Cyperus iria<br />
rice root knot nematode<br />
(damage up to 50%); also<br />
attacks wheat, and<br />
Echinochloa colona<br />
Echinochloa crus-galli,<br />
Portulaca oleracea,<br />
tobacco<br />
wheat, Panicum spp, tomato,<br />
capsicum, etc<br />
Ageratum conyzoides, Croton<br />
lobatus, Cynodon dactylon,<br />
tobacco<br />
attacks tomato and weeds<br />
including Bidens pilosa,<br />
Euphorbia heterophylla,<br />
Galinsoga parviflora<br />
also attacks Cynodon dactylon<br />
E. indica is a moderately good<br />
host of the maize nematode;<br />
has other weed hosts, including<br />
Cyperus iria<br />
Table 4.8.3 Natural enemies of Eleusine indica: fungi and bacteria.<br />
De Waele et al. 1990<br />
Femandez and Ortega<br />
1982<br />
Guo et al. 1984, Holm<br />
et al. 1977<br />
Tedford and Fortnum<br />
1988, Valdez 1968<br />
Rao et al. 1970<br />
Acosta et al. 1986<br />
Lordello et al. 1988<br />
Holm et al. 1977<br />
Femandez and Ortega<br />
1982, Jordaan et al.<br />
1988<br />
Linford and Yap 1940<br />
Species Country Comments References<br />
FUNGI<br />
Bipolaris setariae<br />
(as Drechslera setariae)<br />
Corticium sasakii<br />
Drechslera gigantea<br />
Helminthosporium sp.<br />
Helminthosporium holrnii<br />
Helrninthosporiurn maydis<br />
e<br />
USA (not<br />
recorded in<br />
Australia)<br />
India<br />
Brazil<br />
Thailand<br />
India<br />
China<br />
heavy attack on E. indica,<br />
light on maize, sorghum,<br />
none on dicotyledons<br />
rice, many weeds including<br />
Commelina benghalensis,<br />
Cynodon dactylon,<br />
Firnbristylis miliacea<br />
no hosts other than E. indica<br />
mentioned<br />
also on Echinochloa colona,<br />
Chloris gayana<br />
attacks 2 1 other weeds<br />
including Irnperata cylindrica,<br />
Digitaria ciliaris and<br />
Echinochloa crus-galli<br />
Figliola et al. 1988<br />
Hiremath and<br />
Sulladmath 1985<br />
Roy 1973<br />
Muchovej 1987<br />
Chandrasrikul 1962<br />
Singh and Misra 1978<br />
Wu and Liang 1984<br />
(continued on next page)
92 Biological Control of Weeds: Southeast Asian Prospects<br />
Table 4.8.3 (continued)<br />
Species Country Comments References<br />
Helminthosporium nodulosum<br />
(as Bipolaris nodulosa or<br />
Cochliobolus nodulosus)<br />
Magnaporthe (= Pyricularia)<br />
grisea<br />
Pellicularia rolfsii<br />
Phyllachora eleusines<br />
Pyricularia oryzae<br />
Sclerophthora macrospora<br />
Sclerotiurn rolfsii<br />
Ustilago sp.<br />
Ustilago eleusinis (as<br />
Melanopsichiurn eleusinis)<br />
BACTERIA<br />
Pseudornonas glumae<br />
Pseudornonas plantarii<br />
Africa,<br />
Australia,<br />
India, Japan,<br />
Philippines,<br />
USA<br />
Africa,<br />
Australia,<br />
India, USA,<br />
Georgia<br />
Australia,<br />
India<br />
Africa,<br />
Australia<br />
Brazil, China<br />
India<br />
Australia,<br />
India<br />
China<br />
Africa, Asia<br />
Japan<br />
Japan<br />
also infests maize, Eleusine<br />
coracana, wheat, barley, oats<br />
and weeds including<br />
Dactyloctenium aegyptium;<br />
causes seedling blight leaf<br />
stripe and sooty heads in<br />
E. indica<br />
heavy attack on E. coracana,<br />
Rottboellia cochinchinensis,<br />
light attack on maize<br />
causes wilt disease of E.<br />
coracana and infests many<br />
grasses and dicotyledonous<br />
plants<br />
only recorded on Eleusine and<br />
Eragrostis in Africa; in<br />
Australia only on Eragrostis<br />
attacks rice<br />
attacks maize, wheat, oats,<br />
rice: attacks E. coracana and<br />
many grasses, but not E. indica<br />
in Australia; there may be host<br />
specific strains<br />
attacks many dicotyledonous<br />
crop plants and a wide range<br />
of grasses<br />
smut fungus of Eleusine and<br />
Dactyloctenium, but only on<br />
D. radulans in Australia<br />
an important rice pathogen:<br />
attacks a wide range of weeds<br />
attacks rice, wheat, sorghum,<br />
maize and many weeds<br />
Rachie and Peters<br />
1977, Wapshere<br />
1990b<br />
Chauhan & Verma<br />
198 1, Figliola et al.<br />
1988, Heath et al.<br />
1990, 1992, Shetty et<br />
al. 1985, Valent et al.<br />
1986, Vodianaia et al.<br />
1986, Wapshere<br />
1990b,c<br />
Wapshere 1990b<br />
Parbery 1967,<br />
Ramakrishran 1963<br />
Prabhu et al. 1992,<br />
Teng 1932, Valent et<br />
al. 1986<br />
Rachie and Peters<br />
1977, Ullstrup 1955,<br />
Wapshere 1990b<br />
Reddy 1983, Safeeulla<br />
1976<br />
Mundkur 1939<br />
Simmonds 1966,<br />
Zundel 1953<br />
Miyagawa et al. 1988<br />
Miyagawa et al. 1988
Table 4.8.4 Natural enemies of Ebusine indica: viruses.<br />
Virus Country Other hosts<br />
cereal chlorotic mottle<br />
corn leaf gall<br />
corn stunt<br />
Eleusine mosaic<br />
groundnut rosette<br />
maize dwarf mosaic<br />
maize streak<br />
rice leaf gall<br />
rice orange leaf<br />
rice ragged stunt<br />
rice tungro bacilliform<br />
rice tungro spherical<br />
rice yellow mottle<br />
sugarcane mosaic<br />
sugarcane streak<br />
tungro<br />
wheat rosette<br />
Australia<br />
Philippines<br />
USA<br />
India<br />
Malawi<br />
USA<br />
Nigeria<br />
Philippines<br />
Philippines<br />
China<br />
Philippines<br />
Philippines<br />
Philippines<br />
Kenya<br />
India<br />
Hawaii<br />
Philippines<br />
China<br />
oats, barley, wheat, maize,<br />
E. coracana, Digitaria ciliaris,<br />
Echinochloa colona;<br />
transmitted by Nesoclutha<br />
pallida<br />
maize<br />
several other weeds<br />
maize, sorghum, E. coracana<br />
and many other hosts<br />
groundnut<br />
maize<br />
maize, but not all cultivars<br />
Cicadulina triangula is<br />
the vector<br />
rice<br />
rice<br />
rice, E. indica and 4 other<br />
weeds<br />
rice, Echinochloa<br />
glabrescens, Monochoria<br />
vaginalis, Paspalum distichum<br />
rice, many weeds<br />
rice, many weeds<br />
rice, two grasses<br />
sugarcane<br />
sugarcane<br />
rice<br />
oats, barley, sorghum,<br />
wheat etc. Laodelphax<br />
striatellus is the vector<br />
4.8 Eleusine indica 93<br />
References<br />
Greber 1979<br />
Agati and Calica 1950<br />
Pitre and Boyd 1970<br />
Rao et al. 1965<br />
Adams 1967<br />
Lee 1964<br />
Ekukole et al. 1989,<br />
Rossel et al. 1984<br />
Agati and Calica 1950<br />
Watanakul 1964<br />
Xie et al. 1984<br />
Salamat et al. 1987<br />
Khan et al. 1991<br />
Khan et al. 1991<br />
Okioma et al. 1983<br />
Chona and Rafay 1950<br />
Holm et al. 1977<br />
Watanakul 1964<br />
Zhang et al. 1981<br />
Table 4.8.5 Natural enemies of Eleusine coracana which may prove to have<br />
a limited host range.<br />
Species Country Portion Comments References<br />
attacked<br />
INSECTS<br />
Diptera<br />
CEClDOMYllDAE<br />
Contarinia sp. Uganda inflores- not the same as the<br />
cence sorghum midge Barnes 1946, 1954a,b,<br />
Contarinia sorghicola: 1956, Geering 1953,<br />
the same or a similar Rachie & Peters 1977<br />
species attacks the<br />
common fallow weed<br />
Sorghum verticilliflorum<br />
(continued on next page)
94 Biological Control of Weeds: Southeast Asian Prospects<br />
Table 4.8.5 (continued)<br />
Species Country Portion Comments References<br />
attacked<br />
Hemiptera<br />
APHlDlDAE<br />
Sitobion<br />
(= Macrosiphum)<br />
leelarnaniae<br />
NEMATODES<br />
Heterodera delvii<br />
FUNGI<br />
Melanopsichium<br />
eleusinis(= Ustilago<br />
eleusinis)<br />
India attacks several millets Raychaudhuri 1983<br />
(not in in India including pearl<br />
Australia) millet Pennisetum<br />
glaucum (=P. typhoideum),<br />
also Andropogon vulgare<br />
India root no other hosts Jairajpuri et al. 1979<br />
cysts mentioned<br />
Asia, Africa a smut fungus: only Simmonds, 1966,<br />
from Eleusine and Wapshere 1990c,<br />
Dactylocteniurn: Zundel 1953<br />
tentatively identified<br />
from D. radulans in<br />
Queensland, but not<br />
from E. indica
4.8 Eleusine indica<br />
Map 4.8.2 Distribution of Eleusine indica in Africa (after Phillips 1972)
96 Biological Control of Weeds: Southeast Asian Prospects<br />
Euphorbia heterophylla<br />
(after Barnes and Chan, 1990)
Map 4.9 Euphorbia heterophylla<br />
4.9 Euphorbia heterophylla 97<br />
Euphorbia heterophylla<br />
There are very few records of natural enemies other than fungi attacking Euphorbia<br />
heterophylla and no study has been made in tropical America where it evolved. However,<br />
from the sparse records of insects attacking species of Euphorbia in Brazil it is likely that<br />
adequately host specific insects do occur. Nevertheless E. heterophylla is regarded as an<br />
important weed in southern Brazil.
98 Biological Control of Weeds: Southeast Asian Prospects<br />
4.9 E uphorbia heterophylla L.<br />
(= E. geniculata = E. prunifolia)<br />
Euphorbiaceae<br />
painted spurge, Mexican fire plant; yaa yaang (Thailand)<br />
Rating<br />
+++ Thai<br />
10 ++ Msia<br />
+ Myan, Laos, Camb, Viet, Phil<br />
Indo<br />
Origin<br />
Tropical and sub-tropical America.<br />
Distribution<br />
Widespread as a weed in the tropical and subtropical regions of the world, notably in<br />
Southeast Asia, but apparently not in Kalimantan or Sulawesi (Indonesia) (Soerjani et al.<br />
1986).<br />
Characteristics<br />
An erect annual, up to about 1 m tall; stem cylindrical, hairy; lower leaves alternate;<br />
stems and leaves with milky latex. The simple or lobed leaves are crowded towards the<br />
top of the stem, with a flat, dichotomously-branched, terminal inflorescence of small yel-<br />
low flowers and large leafy bracts, often with a bright red or cream patch at the base. The<br />
inflorescence consists of many small, short-stalked flowers lacking petals but with con-<br />
spicuous glands (Wilson 1981). Reproduction is by seeds which are shed with an explo-<br />
sive mechanism.<br />
Importance<br />
A weed of increasing importance in upland fields of rice and many other crops; also in<br />
wastelands, roadsides, boundaries of coffee plantations; very abundant locally. Seeds per-<br />
sist in the soil until favourable conditions allow germination and rapid growth, giving<br />
rise to large populations of the weed. It is an important weed in 23 tropical countries and<br />
present in at least 37 others. Its rapid growth enables it to compete successfully with<br />
crops, quickly forming a dense canopy over young crop plants. Dense populations of the<br />
weed, with its white sticky latex, may make it impossible to harvest the crop.<br />
The young leaves are sometimes used as a vegetable, but are laxative if too much is<br />
eaten. The plant is said to have caused poisoning in livestock (Wilson 1981).<br />
Natural enemies<br />
Except possibly for Alternaria sp. and Helminthosporium sp. which have not been shown<br />
to be pathogenic to crop plants (Yorinori 1985), there are no records of apparently host
4.9 Euphorbia heterophylla 99<br />
specific organisms attacking Euphorbia heterophylla (Table 4.9.1). However, it is known<br />
that a number of insects do attack it in Brazil, but this observation was incidental to a<br />
study of fungi and none of the insects were identified (E.G. Fontes, pers. comm. 1992).<br />
Although periodic collections were made in Trinidad in the early 1970's, no promising<br />
insects were encountered (Yaseen 1972).<br />
There are few records (19 only) of insects attacking members of the genus<br />
Euphorbia in Brazil (Table 4.9.2) (d'Araujo e Silva et al. 1968a,b), indicating that little<br />
attention has so far been paid to Euphorbia spp. in this region. Six of the insects are<br />
polyphagous and too little is known about the others to arrive at a conclusion. Even if<br />
some are restricted to the Euphorbiaceae, it remains to be determined whether any will<br />
attack either Euphorbia heterophylla or E. hirta.<br />
E. heterophylla is resistant to most herbicides and, in recent years, has become pro-<br />
gressively more important in Brazil, particularly in the southern, soybean-producing<br />
states (Yorinori 1985), which suggests that its insect enemies, if any, may be heavily par-<br />
asitised.<br />
A biological control program has been in progress in Canada since the late 1960's<br />
against Euphorbia cyparissias and E. pseudovirgata, involving the introduction of some<br />
twenty species of insects from Europe. Several species have become established, with<br />
rather localised effects (Julien 1992). It is said that insects are generally unable to attack<br />
Euphorbia species because of the latex that flows freely from any wound and clogs the<br />
mouthparts (Best et al. 1980), but clearly some insects are adapted to deal with this prob-<br />
lem.<br />
The best known economic plant in the Euphorbiaceae is cassava, Manihot esculenta<br />
of South American origin. The insects attacking it there are comparatively well known, a<br />
factor that will aid the evaluation of the specificity of insects attacking Euphorbia spp.<br />
Another species of horticultural importance is poinsettia, Euphorbia pulcherrima.<br />
Table 4.9.1 Natural enemies of Euphorbia heterophylla.<br />
Species Location Other hosts References<br />
INSECTS<br />
Orthoptera<br />
ACRlDlDAE<br />
Poekilocerus<br />
hieroglyphicus<br />
Hemiptera<br />
ALEYRODIDAE<br />
Bemisia tabaci<br />
ALYDIDAE<br />
Leptocorisa acuta<br />
Leptocorisa oratorius<br />
Leptocorisa<br />
solomonensis<br />
Sudan beans, melons Ba-Angood 1977,<br />
Ba-Angood &<br />
Khidir 1975<br />
Thailand, cotton, polyphagous Debrot & Centeno 1985,<br />
Venezuela Nachapong & Mabbett 1979<br />
PNG<br />
PNG<br />
PNG<br />
F. Dori pers. comm. 1993<br />
F. Dori pers. comm. 1993<br />
F. Dori pers. comm. 1993<br />
(continued on next page)
100 Biological Control of Weeds: Southeast Asian Prospects<br />
Table 4.9.1 (continued)<br />
MITES<br />
FUNGI<br />
Species Location Other hosts References<br />
TETRANYCHIDAE<br />
Tetranychus urticae Cuba polyphagous Perez et al. 1987<br />
Alternaria sp.<br />
Amphobotrys ricini<br />
Elsinoe sp.<br />
Helminthosporium sp.<br />
Macrophornina<br />
phaseolinu<br />
Phytophthora<br />
palmivora<br />
Puccinia sp.<br />
Rhizoctonia solani<br />
Sclerotinia<br />
sclerotiorum<br />
Sphaceloma sp.<br />
Uromyces euphorbiae<br />
NEMATODES<br />
Meloidogyne exigua<br />
Meloidogyne javanica<br />
Rotylenchulus<br />
reniformis<br />
Brazil<br />
USA<br />
Burundi<br />
Brazil<br />
India<br />
cassava<br />
Sarawak black pepper Anon 1979<br />
Brazil<br />
Brazil<br />
Brazil<br />
Brazil,<br />
Burundi<br />
Brazil<br />
Brazil<br />
Brazil<br />
Florida<br />
VIRUSES<br />
Euphorbia mosaic Brazil, USA,<br />
Venezuela<br />
cassava<br />
Yorinori 1985<br />
Holcomb et al. 1989<br />
Zeigler & Lozano 1983<br />
Fontes et al. 1992,<br />
Gazziero et al. 1988,<br />
Yorinori 1985<br />
Saxena et al. 198 1<br />
Fontes et al. 1992<br />
Yorinori 1985<br />
Yorinori 1985<br />
Yorinori 1985,<br />
Zeigler & Lozano 1983<br />
Yorinori 1985<br />
coffee, many weeds Luc et al. 1990<br />
Lordello et al. 1988<br />
Inserra et al. 1989,<br />
MacGowan 1989<br />
Debrot & Centeno 1985,<br />
Kim & Flores 1979,<br />
Kim & Fulton 1984,<br />
Yorinori 1985<br />
Table 4.9.2 Insects attacking species of Euphorbia in Brazil (d'Araujo e Silva et al.<br />
1968a,b).<br />
Insect Hosts Feeding habit<br />
Hemiptera<br />
ALEYRODIDAE<br />
Bemisia tabaci<br />
(= B. costa-limai) Euphorbia hirtella, polyphagous<br />
tomato, Mentha arvensis<br />
COCCIDAE<br />
Coccus spp. Euphorbiaceae, Acalypha sp., polyphagous<br />
Aspidosperma ramiflorum,<br />
Cassia sp., Citrus spp.<br />
(continued on next page)
4.9 Euphorbia heterophylla 101<br />
Insect Hosts Feeding habit<br />
Eucalymnatus spp.<br />
Platinglisia noacki<br />
COREIDAE<br />
Chariesterus armatus<br />
TlNGlDAE<br />
Corythuca pellucida<br />
Corythuca socia<br />
Thysanoptera<br />
PHLAEOTHRIPIDAE<br />
Haplothrips gowdeyi<br />
Coleoptera<br />
CHRYSOMELIDAE<br />
Caryedes (= Gibbobruchus)<br />
pickeli<br />
Disonycha argentiniensis<br />
Gibbobruchus polycoccus<br />
CURCULIONIDAE<br />
Sternocoelus sp.<br />
Sternocoelus notaticeps<br />
Lepidoptera<br />
LYMANTRIIDAE<br />
Thagona tibialis<br />
NOCTUIDAE<br />
Spodoptera eridania<br />
NYMPHALIDAE<br />
Didonis biblis<br />
Dynamine artemisia<br />
Episcada pascua<br />
SPHlNGlDAE<br />
Erinnyis oenotrus<br />
Euphorbia capansa, Nerium sp., polyphagous<br />
Caryota sp., Phoenix sp.<br />
Euphorbiaceae, Begonia sp., polyphagous<br />
Eugenia sp., Grevillea robusta,<br />
Ilex sp., kurus sp., Magnolia<br />
pumila, etc.<br />
Euphorbia braziliensis<br />
Euphorbiaceae<br />
Euphorbiaceae<br />
Euphorbia sp., coffee, rice,<br />
Crotolaria sp., PassiJlora sp.,<br />
Buddleia variabilis<br />
Euphorbiaceae<br />
Euphorbia pulcherrima<br />
Euphorbiaceae<br />
Euphorbiaceae<br />
Euphorbiaceae<br />
E. cespitosa, E. ovalifolia,<br />
E. pulcherrima<br />
Euphorbiaceae, many crops<br />
Euphorbiaceae, Tragia volubilis<br />
Euphorbiaceae<br />
Euphorbiaceae<br />
E. ovalifolia<br />
possibly restricted<br />
possibly restricted<br />
possibly restricted<br />
polyphagous<br />
possibly restricted<br />
possibly restricted<br />
possibly restricted<br />
?restricted to Euphorbiaceae<br />
?restricted to Euphorbiaceae<br />
?restricted to Euphorbiaceae<br />
polyphagous<br />
possibly restricted<br />
?restricted to Euphorbiaceae<br />
?restricted to Euphorbiaceae<br />
possibly restricted
102 Biological Control of Weeds: Southeast Asian Prospects<br />
Euphorbia hirta<br />
(after Holm st a/. 1977)
Map 4.10 Euphorbia hirta<br />
4.10 Euphorbia hirta<br />
Euphorbia hirta<br />
There is only one record of a natural enemy attacking Euphorbia hirta in tropical<br />
America where it evolved and only a few of polyphagous species attacking it elsewhere.<br />
A survey in Central America would be necessary to determine what species attack it<br />
there that might be potential biological control agents.
104<br />
Biological Control of Weeds: Southeast Asian Prospects<br />
4.10 Euphorbia hirta L.<br />
(= E. pilulifera)<br />
Euphorbiaceae<br />
garden spurge, asthma plant; mayo (Myanmar), nam nom raatchasee (Thailand)<br />
tuk das khla thom (Cambodia), co sua 16ng (Vietnam), ara tanah, hairy spurge<br />
(Malaysia) gelang susu, gendong ancok (Indonesia), gatas gatas (Philippines)<br />
Rating<br />
++ Thai, Sing, Phil<br />
10 + Laos, Camb, Viet, Msia<br />
w Myan, Indo<br />
Origin<br />
Tropical America.<br />
Distribution<br />
E. hirta is a weed of the tropics and subtropics.<br />
Characteristics<br />
A small, prostrate, hairy annual, 0.15 to 0.3 m tall, with a tap root; stems much branched<br />
from the base, often reddish, bearing brownish stiff hairs and having milky sap; leaves,<br />
hairy, opposite, sometimes purple-blotched and with toothed margins; flowers unisexual;<br />
reproduction by seeds 0.5 to 1 mm long.<br />
Importance<br />
E. hirta grows well in sunny to lightly shaded cultivated lands, gardens, lawns, waste<br />
areas and run down grasslands. It is an early coloniser of bare ground especially under<br />
damp or irrigated conditions. It flowers all year round in Southeast Asia producing up to<br />
3000 seeds per plant. When the seed pods mature they explode to disperse the seeds. Its<br />
prostrate habit enables it to tolerate mowing and it can be important in lawns. It has been<br />
reported from 47 countries as a weed in many crops, including citrus, cotton, groundnuts,<br />
maize, pineapples, rice, sorghum, sugarcane, tea and vegetables. Moody (1 989) records it<br />
as being more widespread in rice than Euphorbia heterophylla.<br />
E. hirta is sometimes used in medicines in Fiji, Malaysia, Indonesia, the Philippines<br />
and Brazil-the leaves and latex against intestinal diseases, ulcers and bronchitis, and the<br />
latex for conjunctivitis. It may have slightly poisonous properties and is useless as fodder<br />
for livestock.<br />
Natural enemies<br />
In view of its common occurrence in the tropical and subtropical belt of the world, it is<br />
surprising that there are so few records of natural enemies attacking it, and those that do<br />
are highly polyphagous (Table 4.10.1). A survey in Central America would be necessary<br />
to learn more about its natural enemies that might have potential for biological control.
Table 4.10.1 Natural enemies of Euphorbia hirta.<br />
4.10 Euphorbia hirta 105<br />
Species Location Other hosts References<br />
INSECTS<br />
Orthoptera<br />
ACRlDlDAE<br />
Chrotogonus<br />
trachypterus<br />
Hemiptera<br />
APHlDlDAE<br />
Aphis craccivora<br />
Aphis gossypii<br />
ALEYRODIDAE<br />
Bemisia tabaci<br />
DELPHACIDAE<br />
Tarophagus proserpina<br />
LYGAEIDAE<br />
Nysius inconspicuus<br />
PSEUDOCOCCIDAE<br />
Ferrisia virgata<br />
Thysanoptera<br />
THRlPlDAE<br />
Haplothrips euphorbiae<br />
Diptera<br />
AGROMYZIDAE<br />
Liriomyza bryoniae<br />
Liriomyza strigata<br />
Lepidoptera<br />
NOCTUI DAE<br />
Achaea janata<br />
FUNGI<br />
Aecidium tithymali<br />
Amphobotrys ricini<br />
Cylindrocladium<br />
quinqueseptatum<br />
Phytophthora palmivora<br />
PROTOZOA<br />
Phytomonas sp.<br />
NEMATODES<br />
Meloidogyne incognita<br />
India<br />
Nigeria,<br />
Uganda<br />
India<br />
India<br />
Philippines<br />
India<br />
India<br />
India<br />
Europe<br />
W. Europe,<br />
USSR<br />
Indonesia<br />
Thailand<br />
USA<br />
India<br />
Sarawak<br />
Venezuela<br />
Hawaii<br />
polyphagous Chandra et al. 1983<br />
poly phagous, Booker 1964, Davies 1972,<br />
a virus transmitter Ofuya 1988<br />
polyphagous Jeritta & David 1986<br />
polyphagous, Jeyarajan et al. 1988<br />
a virus transmitter<br />
polyphagous Duatin & Pedro 1986<br />
polyphagous Thangavelu 1978<br />
poly phagous Jeritta & David 1986<br />
possibly host restricted Jeritta & David 1986<br />
highly polyphagous Spencer 1973, 1990<br />
highly polyphagous Spencer 1973, 1990<br />
polyphagous Kalshoven 198 1<br />
Puckdeedindan 1966<br />
Holcomb et al. 1989<br />
Sulochana et al. 1982<br />
black pepper Anon 1979<br />
Barreto 1982<br />
Valdez 1968<br />
(continued on next page)
106 Biological Control of Weeds: Southeast Asian Prospects<br />
Table 4.10.1 (continued)<br />
Species Location Other hosts References<br />
Meloidogyne javanica India Dahiya et al. 1988<br />
Radopholus sirnilis Zimbabwe polyphagous Martin et al. 1969<br />
Rotylenchulus reniforrnis Hawaii, USA Linford & Yap 1940,<br />
Inserra et al. 1989<br />
VIRUSES<br />
groundnut rosette Hawaii, Nigeria, Adams 1967,<br />
Uganda Booker 1964,<br />
Davies 1972<br />
hibiscus yellow India Jeyarajan et al. 1988<br />
vein mosaic<br />
tapioca mosaic India Jeyarajan et al. 1988<br />
tobacco leaf curl Hawaii Holm et al. 1977<br />
tomato leaf curl India Jeyarajan et al. 1988<br />
urd bean yellow mosaic India Jeyaragan et al. 1988
108 Biological Control of Weeds: Southeast Asian Prospects<br />
Fimbristylis miliacea<br />
(after Holm et a/. 1977)
Map 4.11 Fimbristylis miliacea<br />
4.1 1 Fimbristylis miliacea 109<br />
Fimbristylis miliacea<br />
Very few natural enemies of Fimbristylis miliacea are known and it would be necessary<br />
to carry out a survey in tropical America before it would be possible to evaluate the<br />
prospects for its biological control.
110 Biological Control of Weeds: Southeast Asian Prospects<br />
4.11 Fimbristylis miliacea (L.) Vahl<br />
(= l? littoralis)<br />
Cyperaceae<br />
lesser fimbristylis, grass-like fimbristylis; m6nhnyin (Myanmar), kak phrek kdam,<br />
smao (Cambodia), rumput bukit, rumput tahi berbau (Malaysia), agor (Thailand),<br />
ba bawagan (Indonesia)<br />
Rating<br />
+++ Myan, Camb, Msia, Brun, Indo<br />
23 ++ Viet, Sing, Phil<br />
. Thai, Laos<br />
Origin<br />
Tropical America.<br />
Distribution<br />
Central America, West Africa, Asia and Southeast Asia to northern Australia.<br />
Characteristics<br />
An erect annual or perennial sedge, growing up to 0.9 m; flower stems 4 or 5 angled,<br />
leaves two-ranked, threadlike, stiff and half as long as flower stems; inflorescence a dif-<br />
fuse compound umbel.<br />
Importance<br />
F. miliacea thrives in damp, open waste places, competing actively with other vegetation<br />
following germination during dry periods or shallow water conditions. A layer of water<br />
15 cm deep suppresses germination. Seedlings emerge during the entire growing period<br />
of rice with which it competes actively. It is a troublesome weed in 21 countries. In<br />
Malaysia it is the first sedge to emerge after rice has been transplanted and the first to<br />
recover after ploughing. In the Philippines it flowers all year, plants each producing up to<br />
10000 seeds. In many places there is no seed dormancy.<br />
F. miliacea is one of the most serious and widespread weeds of rice and is also<br />
reported from taro (Hawaii), bananas (Taiwan), abaca (Philippines), maize, sugarcane<br />
(Indonesia, Taiwan) and sorghum (Malaysia).<br />
F. miliacea is eaten by cattle, but the seeds are mostly undigested and germinate<br />
near the dung.<br />
~atu'ral enemies<br />
So little is known about its natural enemies (Table 4.1 1.1) that it is not possible to evalu-<br />
ate the prospects for biological control. A survey is necessary in tropical America.
Table 4.11.1 Natural enemies of Fimbrisstylis miliacea.<br />
4.1 1 Fimbristylis miliacea 111<br />
Species Location Other hosts References<br />
INSECTS<br />
Hemiptera<br />
PENTATOMIDAE<br />
Scotinophara Philippines rice, Cornrnelina Barrion & Litsinger 1987<br />
latiuscula benghalensis,<br />
Echinochloa crus-galli<br />
Lepidoptera<br />
Creatonoros gangis Philippines rice, many weeds Catindig et al. 1993<br />
(= Amsacra)<br />
FUNGI<br />
Corticum sasakii India Commelina Roy 1973<br />
benghalensis,<br />
Cynodon dactylon,<br />
Eleusine indica and<br />
other grasses<br />
NEMATODES<br />
Criconemella onoensis rice, many weeds Luc et al. 1990<br />
Hirschmaniella spp. rice, maize, sugarcane, Luc et al. 1990<br />
many weeds<br />
Meloidogyne India rice, many weeds Luc et al. 1990,<br />
graminicola Rao et al. 1970<br />
Meloidogyne oryzae Surinam rice, plantain, wheat, Maas et al. 1978<br />
potato, tomato<br />
Rotylenchulus Trinidad very polyphagous Singh 1974<br />
reniformis
112 Biological Control of Weeds: Southeast Asian Prospects<br />
Marsilea minu fa<br />
(after Soerjani el a/. 1987)
Map 4.12 Marsilea minuta<br />
4.1 2 Marsilea minuta 113<br />
Marsilea minuta<br />
Marsilea minuta, water clover, is thought to be of tropical African origin, but no account<br />
of its natural enemies there is available. A survey would thus be required to evaluate the<br />
prospects for its biological control.
114 Biological Control of Weeds: Southeast Asian Prospects<br />
4.12 Marsilea minuta L.<br />
(=M. crenata)<br />
Marsileaceae<br />
water clover, clover fern, pepperwort; pak vaen (Laos), chuntul phnom<br />
(Cambodia), semanggi (Indonesia), phak waen (Thailand), tapah itik (Malaysia)<br />
paang itik (Philippines).<br />
Rating<br />
+++ Msia<br />
12 ++ Indo, Phil<br />
+ Myan, Thai, Laos, Camb, Viet<br />
Origin<br />
Africa or possibly tropical Asia (Jacobsen 1983). It consists of a complex of strains<br />
including a diploid (n = 20) and a sterile triploid (2n = 60) (Tryon and Tryon 1982).<br />
Distribution<br />
Marsilea minuta is widespread over most of the African continent and it is pantropical in<br />
Asia. The Marsileaceae contains about 65 species, of which 16 occur in Africa and, of<br />
these, M. minuta is amongst the most widespread (Jacobsen 1983).<br />
Characteristics<br />
A very variable, perennial water fern of aquatic or marshy sites. Its stems are creeping<br />
rhizomes rooted in the mud. Leaves are clover-like, with four leaflets borne on a petiole 2<br />
to 30 cm long. Leaflets have fan-shaped, repeatedly bifurcating veins and normally float<br />
on the water surface. Sporocarps occur near the base of the petioles and usually occur<br />
under the mud or water surface. Reproduction is by spores or rhizomes.<br />
Importance<br />
Although M. minuta has a rating of 12 and is widely reported as a weed in Southeast<br />
Asia, there are surprisingly few references to it as a weed in the literature except for<br />
those dealing with chemical control. In Thailand it is common in rice fields and along<br />
canals and other waterways. It is one of the seven most important weeds in the Muda<br />
area of Malaysia (Itoh 1991a). It is one of the more important emergent weeds in shallow<br />
water rice fields in the central lowlands in Vietnam (Nguyen Van Vuong 1973) and in the<br />
lowland area of Vientiane in Laos (Sisounthone and Sisombat 1973). In Indonesia its<br />
vegetative growth and reproduction is very rapid. It can grow under water and, after<br />
weeding, rapidly re-establishes itself unless well buried in the soil. It is an effective com-<br />
petitor for nutrients, particularly in the early part of the growth period after transplanta-<br />
tion of rice seedlings when M. minuta rapidly covers the ground surface. In the<br />
Philippines it caused 19% crop loss when sown together with rice (Suriapermana 1977).<br />
The young leaves of water clover are sometimes eaten as a vegetable in Indonesia.
Natural enemies<br />
4.1 2 Marsilea minuta 115<br />
Very little information concerning natural enemies emerged from computer-aided search-<br />
es of the literature on Marsilea minuta, which also included searches of its synonyms:<br />
M. crenata and M. erosus in Asia and M. difisa, M. perrieriana and M. senegalensis in<br />
Africa (Table 4.12.1). In Africa Marsilea minuta appears to be regarded, at most, as a<br />
minor weed. This is possibly due to the fact that rice is far less important there than in<br />
Asia, or it may be due to the presence of effective natural enemies in Africa, although<br />
these have not yet been reported.<br />
In the Philippines, larvae of the ephydrid flies Notiphila Eatigenis and N. similis are<br />
common on emergent M. minuta and damage its stems. Their eggs are usually laid on the<br />
stems and serve as alternative hosts of Trichogramma wasps attacking rice stem borers<br />
(Barrion and Litsinger 1986). The contents of upwards of 90% of the sporocarps from<br />
M. minuta growing under terrestrial conditions in northwestern India were destroyed by<br />
larvae of the weevil Echinocnemus. The larval and pupal stages are completed in 40 to 45<br />
days (Loyal and Kumar 1977). In Indonesia the case-forming larvae of the widely distrib-<br />
uted pyralid moth Elophila (= Nymphula) responsalis attacked M. minuta and several<br />
other aquatic plants including Salvinia spp., Lemna purpusilla, L. polyrhiza, Monochoria<br />
vaginalis, Azolla pinnata and Pistia stratiotes. However, tests have shown that it will not<br />
feed on rice. The development period of Elophila responsalis ranged from 42 to 56 days.<br />
It was attacked by a pupal parasitoid (Tetrastichus sp.) and a larval coleopteran predator<br />
(Handayani and Syed 1976, Sankaran and Rao 1972, Subagyo 1975). Elophila respon-<br />
salis occurs also in India, Sri Lanka, Myanmar, Japan and Australia.<br />
Comment<br />
A survey for natural enemies attacking M. minuta in Africa is required before the<br />
prospects for its biological control in Southeast Asia can be evaluated.<br />
Table 4.12.1 Natural enemies of Marsilea minuta.<br />
Species Location References<br />
INSECTS<br />
Coleoptera<br />
CURCULIONIDAE<br />
Echinochnemus sp.<br />
Diptera<br />
EPHYDRIDAE<br />
Notiphila latigenis<br />
Notiphila similis<br />
India Loyal & Kumar 1977<br />
Philippines Barrion & Litsinger 1986<br />
Philippines Barrion & Litsinger 1986<br />
(continued on next page)
116 Biological Control of Weeds: Southeast Asian Prospects<br />
Table 4.12.1 (continued)<br />
Species Location References<br />
Lepidoptera<br />
PY RALl DAE<br />
Elophila (= Nymphula)<br />
responsalis<br />
FUNGI<br />
VIRUS<br />
Alternaria sp.<br />
Cercospora rnarsileae<br />
Phaeotrichoconis crotalariae<br />
Pistia virus<br />
India, Indonesia<br />
India<br />
India<br />
India<br />
India<br />
Handayani & Syed 1976,<br />
Sankaran & Rao 1972,<br />
Subagyo 1975<br />
Menon & Ponnappa 1964<br />
Patil 1975<br />
Menon & Ponnappa 1964<br />
Menon & Ponnappa 1964
118 Biological Control of Weeds: Southeast Asian Prospects<br />
Melastoma malabathricum<br />
(after Soerjani et a/. 1987)
Map 4.13 Melastoma malabathricum<br />
4.1 3 Melastoma malabathricum 119<br />
Melastoma malabathricum<br />
M. rnalabathricum is a perennial shrub which probably originated in Southeast Asia or<br />
neighbouring areas, including Irian Jaya, Papua New Guinea and northern Australia, a<br />
region where it is regarded as being of little importance. A survey in this region would<br />
reveal whether there are promising natural enemies for biological control.
120 Biological Control of Weeds: Southeast Asian Prospects<br />
4.13 Melastoma malabathricum L.<br />
(= Melastoma affine)<br />
Melastomataceae<br />
melastoma, Indian rhododendron, Straits rhododendron; senduduk (Malaysia)<br />
Rating<br />
+++ Msia, Brun<br />
13 ++ Sing<br />
+ Thai, Laos, Viet, Indo, Phil<br />
Camb<br />
Origin<br />
Asia, Papua New Guinea, Australia.<br />
Characteristics<br />
M. malabathricum is a perennial shrub growing to 2 m high; its stems are reddish with<br />
rough upwardly pointing scales, the leaves are tapered to both ends, are rough to touch<br />
and have three distinct ribs. The flowers, which are clustered at the ends of twigs, are<br />
pinkish to light violet. The fruit is a berry-like capsule covered with scales.<br />
Importance<br />
Melastoma is common in abandoned clearings, on waste ground and in cultivated lands.<br />
In addition to its importance in Southeast Asia, it is a principal weed of rubber in West<br />
Africa. It is said to make good firewood. The sweetish black seeds are eaten and chewed<br />
leaves are used for burns and against amoebic dysentery. The fruits host a fruit fly<br />
species in the Bactrocera dorsalis complex which does not attack commercially impor-<br />
tant fruits (R.A.I. Drew, pers. comm.).<br />
Natural enemies<br />
Krauss (1 965) surveyed the natural enemies of species of Melastoma, including M. mala-<br />
bathricum, in various countries of Southeast Asia and islands of the western Pacific.<br />
Twenty six insect species were found on M. malabathricum (Table 4.13.1) and a further<br />
34 species on other melastomas. It is very likely that some of the 34 species will also<br />
attack M. malabathricum and, indeed, the leaf rolling pyralid moth Ategumia fatualis<br />
does so. After specificity tests Ategumia fatualis was liberated in Hawaii and Kauai in<br />
1958 and became established, although it did not become sufficiently abundant to pro-<br />
vide effective control (Table 4.13.2) (Krauss 1965).<br />
Another leaf-rolling pyralid Ategumia adipalis was liberated in 1965, and was<br />
reported to have become established (Davis and Chong 1969), but at low population lev-<br />
els. Next a noctuid moth Selca brunella was introduced to Kauai and Hawaii from<br />
Malaysia and Singapore in 1964 and was recovered the next year. The larvae feed avidly
4.13 Melastoma malabathricum 121<br />
in flower buds, bore into terminal stems and eat leaves. In heavily infested localities con-<br />
siderable dieback has resulted, at places flowering was prevented and in others up to 50%<br />
of fruits were destroyed by larvae. Larvae have been found recently on Tiboochina urvil-<br />
leana and Heterocentron subtriplinenium (both Melastomataceae) in Hawaii (C.J. Davis<br />
pers. comm. 1993). A braconid wasp Meteorus sp. attacks S. brunella larvae (Davis<br />
1970, Davis and Chong 1969).<br />
An unidentified grasshopper and an unidentified lepidopterous larva attack M. mal-<br />
abathricum in Thailand but not the chrysomelid beetle Altica cyanea which is present<br />
there and attacks it in Indonesia and Malaysia (Napompeth 1982).<br />
Comment<br />
Although a number of insects are known to attack M. malabathricum in Southeast Asia<br />
(and especially in Malaysia), they clearly do not reduce its status to the level required and<br />
thus are of limited value for classical biological control in that region. However, if as<br />
postulated, the area of origin includes Papua New Guinea, (Irian Jaya) and Australia it is<br />
possible that there may be useful natural enemies in the region that do not occur in coun-<br />
tries to the north and west. Certainly, Melastoma is not listed as an important weed in<br />
Papua New Guinea.<br />
Table 4.13.1 Natural enemies of Melastoma malabathricum.<br />
Species Country Food References<br />
INSECTS<br />
Hemiptera<br />
APHlDlDAE<br />
Aphis sp.<br />
CICADELLIDAE<br />
Tettigella<br />
(= Tettigoniella) sp.<br />
COCCI DAE<br />
Rastrococcus sp.<br />
MEMBRACIDAE<br />
Gargara sp.<br />
kptocentrus taurus<br />
Nilaurama minutispina<br />
Sipylus dilatatum<br />
Sipylus sp.<br />
Tricentrus sp.<br />
MlRlDAE<br />
Helopeltis antonii<br />
Hyalopeplus vitripennis<br />
RlCANllDAE<br />
- Pochazia antica<br />
Singapore leaf<br />
Malaysia leaf<br />
Malaysia<br />
Malaysia<br />
Malaysia<br />
Malaysia<br />
Malaysia<br />
Malaysia<br />
Malaysia<br />
Singapore<br />
Indonesia<br />
Malaysia<br />
Malaysia<br />
leaf<br />
branch; also on<br />
Melastoma<br />
polyanthum<br />
branch<br />
branch<br />
branch<br />
leaf<br />
leaf<br />
Krauss 1965<br />
Krauss 1965<br />
Krauss 1965<br />
Krauss 1965<br />
Krauss 1965<br />
Krauss 1965<br />
Krauss 1965<br />
Krauss 1965<br />
Krauss 1965<br />
Krauss 1965<br />
Soerjani et al. 1987<br />
Krauss 1965<br />
Krauss 1965<br />
(continued on next page)
122 Biological Control of Weeds: Southeast Asian Prospects<br />
Table 4.13.1 (continued)<br />
Species Country Food References<br />
Coleoptera<br />
CHRYSOMELIDAE<br />
Altica cyanea Indonesia,<br />
CURCULIONIDAE<br />
Alcidodes sp.<br />
Ceutorhynchus sp.<br />
Cryptorhynchus sp.<br />
Imerodes (?) sp.<br />
Malaysia<br />
Malaysia<br />
Malaysia<br />
Malaysia<br />
Singapore<br />
Diptera<br />
TEPHRITIDAE<br />
Bactrocera dorsalis<br />
(= B. pedestris) Malaysia,<br />
Singapore,<br />
Sri Lanka<br />
Lepidoptera<br />
ARCTllDAE<br />
Species of Lithosiinae<br />
GELECHllDAE<br />
Idiophantis sp.<br />
HYPONOMEUTIDAE<br />
Argyresthia leuculias<br />
LYMANTRIIDAE<br />
?Species<br />
NOCTUIDAE<br />
Autoba (= Eublemma)<br />
versicolor<br />
Selca brunella<br />
PY RALl DAE<br />
Agrotera basinotata<br />
Ategumia adipalis<br />
Ategumia fatualis<br />
TORTRICIDAE<br />
Archips rnicaceana<br />
shoot<br />
flowers<br />
flowers<br />
flowers<br />
Malaysia fruit<br />
Malaysia<br />
Malaysia<br />
Malaysia<br />
Malaysia<br />
Malaysia,<br />
Singapore<br />
Malaysia<br />
Malaysia,<br />
Singapore<br />
Philippines<br />
Kamarudin & Shah<br />
1978, Napompeth 1982<br />
Krauss 1965<br />
Krauss 1965<br />
Krauss 1965<br />
Krauss 1965<br />
fruit Krauss 1965<br />
fruit<br />
fruit<br />
flower<br />
leaf<br />
leaf, twig, fruit<br />
leaf<br />
leaf<br />
Krauss 1965<br />
Krauss 1965<br />
Krauss 1965<br />
Krauss 1965<br />
Krauss 1965<br />
Julien 1992, Krauss 1965<br />
Krauss 1965<br />
Julien 1992, Krauss 1965<br />
leaf Krauss 1965<br />
Malaysia leaf Krauss 1965<br />
FUNGI<br />
Phytophthora palmivora Sarawak black pepper Anon 1979
4.13 Melastoma malabathricum 123<br />
Table 4.13.2 Introductions to Hawaii for the biological control of Melastoma<br />
malabathricum.<br />
Species Source Liberated Established References<br />
INSECT<br />
Lepidoptera<br />
NOLIDAE<br />
Selca brunella<br />
PYRALl DAE<br />
Ategumia adipalis<br />
Ategumia fatualis<br />
Malaysia, 1965 +<br />
Singapore<br />
Malaysia, 1965 +<br />
Singapore<br />
Philippines 1958 +<br />
Davis 1960, Davis &<br />
Chong 1968, Davis &<br />
Krauss 1962, 1966,<br />
1967, Julien 1992,<br />
Krauss 1965<br />
Davis & Chong 1969,<br />
Julien 1992<br />
Davis & Krauss 1966,<br />
Julien 1992
124 Biological Control of Weeds: Southeast Asian Prospects<br />
Mikania micrantha<br />
(after Holm et a/. 1977)
Map 4.14 Mikania micrantha<br />
4.1 4 Mikania micrantha 125<br />
Mikania micrantha<br />
Mikania micrantha is a perennial vine, native to Central and South America.<br />
A number of very promising, and probably specific, natural enemies are known in<br />
Central and South America where M. micrantha is not regarded as a weed. One of these,<br />
a thrips Liothrips mikaniae has been released in Malaysia and the Solomon Is, but<br />
extremely high predation is believed to have prevented its establishment. A bug<br />
Teleonemia sp., several chrysomelid beetles and an eriophyid mite Acalitus sp. warrant<br />
serious consideration. A number of other natural enemies, whose specificity has not yet<br />
been adequately investigated, also attack M. micrantha.<br />
In spite of the lack of success with the thrips, M. micrantha appears to be a prime<br />
target for the introduction of one or more of the other organisms that attack it in its area<br />
of origin.
126 Biological Control of Weeds: Southeast Asian Prospects<br />
4.14 Mikania micrantha Kunth<br />
Asteraceae<br />
mile-a-minute weed; cheroma, ulam tikus (Malaysia), sembung rambat<br />
(Indonesia).<br />
This chapter updates that in Waterhouse and Norris (1987), with special reference to<br />
Southeast Asia.<br />
Rating<br />
+++ Msia, Sing<br />
11 ++ Brun, Indo<br />
+ Phil<br />
Thai<br />
Origin<br />
The weedy species in Southeast Asia and the Pacific is M. micrantha from Central and<br />
South America and not the North American M. scandens or the Old World M. cordata<br />
(Parker 1 972).<br />
Distribution<br />
M. micrantha belongs to a genus containing about 250 species of mostly Central and<br />
South American origin. In addition to its native distribution in tropical America, it has<br />
spread to Mauritius, India, Sri Lanka, Bangladesh and Southeast Asia (as above). It<br />
occurs widely as a weed in the Pacific, including Papua New Guinea (Waterhouse and<br />
Norris 1987), but is not yet present in Australia. It was introduced from Paraguay to<br />
Bogor Botanic Gardens (Indonesia) in 1949 and, in 1956, was used as a soil cover in rub-<br />
ber: it has since spread throughout Indonesia (Soerjani et al. 1987).<br />
Characteristics<br />
Mikania micrantha is an extremely fast growing, sprawling, perennial vine, with oppo-<br />
site, heart-shaped leaves, longitudinally ribbed, branched and hairless stems and numer-<br />
ous small heads of densely clustered white flowers. It creeps and twines, roots readily at<br />
the nodes and produces abundant small (2 mm long) black seeds bearing a terminal tuft<br />
of white bristles that aid wind dispersal.<br />
In its natural habitat in tropical America, M. micrantha is usually found in disturbed<br />
situations. It seldom occurs on poor soils and is most commonly found in damp or<br />
swampy places. Typical sites in South America are roadsides in wet forest and the edges<br />
of freshwater swamps. Flowering occurs mainly in the dry season and only in sunny situ-<br />
ations (Cock 1982a).<br />
Importance<br />
With its rapid growth, ready rooting at nodes, smothering habit and prolific seed produc-<br />
tion, M. micrantha rapidly colonises disturbed habitats, retarding, by competition and
4.14 Mikania micrantha 127<br />
through plant inhibitors that it elaborates (Wong 1964), the growth of crops or natural<br />
vegetation. In comparison with a nitrogen-fixing legume, it is of restricted value in the<br />
role of a cover crop. For example, in Malaysia the girth of rubber trees was 27% less<br />
with a cover of M. micrantha than of a legume and the yield over the first 32 months of<br />
production was 27 to 29% less (Teoh et al. 1985). In many parts of Southeast Asia it is a<br />
serious pest of plantation crops (oil palm, coconut, cocoa, tea, rubber, teak). Its climbing<br />
habit enables it to reach and then dominate the crowns of bushes or trees up to 10m high,<br />
where it is difficult to attack either mechanically or chemically without risk of damaging<br />
the crop (Parker 1972). It recovers rapidly from slashing. It is eaten by cattle, but is less<br />
valuable as fodder than many of the pasture plants it is able to smother. Nevertheless, it is<br />
viewed by some as being useful to control soil erosion, to serve as a mulch when cut, and<br />
as being preferable to many alternative plants that might occupy the space vacated by its<br />
control. In its native habitat it is seldom a weed.<br />
Natural enemies<br />
MAJOR SPECIES<br />
TROPICAL AND SOUTH AMERICA<br />
Valuable information is available on 9 major and 22 minor natural enemies of M. micran-<br />
tha in its native region as a result of studies by Cock (1982a,b) and Freitas (1991).<br />
Several of the major natural enemies are reported to be promising biological control<br />
agents (Table 4.14.1) and all these, and probably some of those less extensively studied<br />
(Table 4.14.2), are worthy of serious consideration. Details of their biology and possible<br />
relevance for biological control are summarised below.<br />
Table 4.14.1 Major natural enemies of Mikania micrantha in its native range in<br />
Central and South America (after Cock 1982a).<br />
INSECTS<br />
Thysanoptera<br />
PHLAEOTHRIPIDAE<br />
Liothrips rnikaniae<br />
Hemiptera<br />
TlNGlDAE<br />
Teleonernia sp. or spp. nr prolixa<br />
Coleoptera<br />
CHRYSOMELIDAE<br />
Desmogramma conjuncta<br />
Echorna rnarginata<br />
Echorna quadristillata<br />
Physimerus pygrnaeus<br />
APlONlDAE<br />
Apion luteirostre<br />
CURCULIONIDAE<br />
' Pseudoderelomus baridiiformis<br />
MITE<br />
ERlOPHYlDAE<br />
Acalitus sp.
128 Biological Control of Weeds: Southeast Asian Prospects<br />
Table 4.14.2 Additional natural enemies of Mikania micrantha, not known to be<br />
polyphagous, from Central and South America (Cock 1982a, Freitas 199 1 ).<br />
Species Distribution Feeding location Host range<br />
INSECTS<br />
Hemiptera<br />
LYGAEIDAE<br />
Xyonysius basalis<br />
(= X. inaequalis<br />
= X. sp. nr ementitus)<br />
MEMBRACIDAE<br />
Entylia sinuata<br />
Entylia sp.<br />
Trinidad, inflorescence<br />
Venezuela<br />
Colombia, stems and leaves<br />
Costa Rica<br />
Trinidad, Peru, stems and leaves<br />
Venezuela<br />
Ecuador<br />
Trinidad, stems and young<br />
Probably other Mikania<br />
spp.or other Asteraceae<br />
M. cordifolia<br />
M. cordifolia<br />
Micrutalis binaria M. vitifolia and<br />
Colombia leaves<br />
M. trinitaria<br />
MlRlDAE<br />
Pycnoderes incurvus<br />
TlNGlDAE<br />
Colombia,<br />
Ecuador,<br />
Costa Rica<br />
leaves<br />
probably specific<br />
Leptocysta sexnebulosa Venezuela<br />
Colombia. Peru<br />
mature leaves M. cordifolia<br />
Coleoptera<br />
CHLAMlSlDAE<br />
Exema complicata Trinidad, Peru, leaves<br />
Costa Rica,<br />
Colombia<br />
CHRYSOMELIDAE<br />
Longitarsus sp.<br />
nr amazonus<br />
Sceloenopla sp.<br />
Colombia, Peru leaves<br />
Trinidad leaves<br />
Diptera<br />
AGROMYZIDAE<br />
Calycomyza mikaniae Trinidad, leaf miner<br />
Colombia,<br />
CEClDOMYllDAE<br />
Neolasioptera sp.<br />
TEPHRlTlDAE<br />
Xanthaciura insecta<br />
Costa Rica<br />
Mikania spp., Chromolaena<br />
odorata and possibly<br />
other Asteraceae<br />
probably specific<br />
M. trinitaria<br />
Mikania spp. and possibly<br />
related Asteraceae<br />
Trinidad, Colombia flowers M. cordifolia<br />
Trinidad flower head<br />
Lepidoptera<br />
GELECHllDAE<br />
Onebala tegulella Trinidad,<br />
Costa Rica<br />
leaf roller<br />
Recurvaria sp. Trinidad flowers<br />
various Asteraceae<br />
M. vitifolia<br />
various Asteraceae<br />
(continued on next page)
4.14 Mikania micrantha 129<br />
Species Distribution Feeding location Host range<br />
GEOMETRIDAE<br />
Chloropteryx sp.<br />
Eupithecia sp.<br />
LYCAENIDAE<br />
Thereus<br />
(= Thecla) palegon<br />
NYMPHALIDAE<br />
Tegosa claudina<br />
(= Tegosa similis)<br />
PTEROPHORIDAE<br />
Adaina bipuncta<br />
PY RALl DAE<br />
Lamprosema distincta<br />
TORTRICIDAE<br />
Lobesia (= Polychrosis)<br />
?carduana<br />
Phalonidia<br />
multistrigata<br />
Trinidad<br />
Trinidad<br />
Trinidad<br />
Brazil, Trinidad,<br />
Colombia<br />
Trinidad<br />
Trinidad, Panama,<br />
Costa Rica<br />
Trinidad<br />
Trinidad<br />
flowers<br />
flowers<br />
flowers<br />
leaves<br />
flowers<br />
leaf roller<br />
flowers<br />
flowers<br />
various Asteraceae<br />
various Asteraceae<br />
various Asteraceae<br />
possibly specific<br />
various Asteraceae<br />
possibly specific<br />
various Asteraceae<br />
various Asteraceae<br />
Acalitus sp. Acarina: Eriophyidae<br />
Feeding on the leaves by this eriophyid mite causes the formation of raised patches<br />
(erinea) in which the mites and their immature stages congregate. In Venezuela the patches<br />
mostly protrude on the undersurface of the leaf, whereas elsewhere they are mostly on<br />
the uppersurface, which may indicate taxonomic differences. At low mite densities the<br />
small number of erineum patches appear to have little effect on the growth and vigour of<br />
the plant. However, in dense infestations, erineum patches cover all the young leaves and<br />
spread into the flower heads, resulting in shortened internodes and reduced flowering.<br />
Plant vigour is significantly reduced.<br />
Eriophyid mites are usually restricted to a single plant species. Although erineum<br />
patches occur widely on M. micrantha, they were not seen on any other species of<br />
Mikania encountered in Cock's (1 982a) studies, suggesting a high degree of specificity.<br />
Predatory or scavenger mites occur quite commonly in and around the erineum patches.<br />
If field specificity trials with potted plants of closely related Asteraceae placed among<br />
heavily infested M. micrantha prove negative, this mite would be a promising biological<br />
control agent. Similar mites on Lantana in South America appear to discourage insect<br />
attack (K.L.S. Harley pers. comm.).<br />
Apion luteirostre Coleoptera: Apionidae<br />
Eggs of this weevil are laid in unopened host flower heads. Larvae feed initially on the<br />
petals, stigma and stamens and, later, destroy the developing seeds. They pupate in the<br />
flower head. The adults make small holes in young leaves.<br />
rn extensive field studies A. luteirostre larvae were not recorded from Chromolaena<br />
odorata, although they were found on M. micrantha and M. vitifolia, but not on M. cordifolia.<br />
Starvation tests using adults resulted in their feeding on five species of Mikania and<br />
on Bidens pilosa, but not on Chromolaena odorata.
130 Biological Control of Weeds: Southeast Asian Prospects<br />
Larvae of Apion luteirostre are attacked by the non-specific eulophid parasitoid<br />
Horismenus? aeneicollis and the pteromalid Zatropis sp. A number of Apion species have<br />
been used successfully in biological control of weeds programs (e.g. Emex australis and<br />
E. spinosa). Further host specificity trials are needed to evaluate the potential value of<br />
A. luteirostre.<br />
Desmogramma conjuncta Coleoptera: Chrysomelidae<br />
This chrysomelid beetle occurs widely, but at low density, on M. micrantha in Central<br />
and South America and a related species D. bigaria occurs on M. micrantha in<br />
Venezuela.<br />
Eggs are laid on the host leaves on which the larvae feed. Pupation occurs in the<br />
soil. No field records are available of feeding on plants other than M. micrantha and, in<br />
limited multiple choice tests, adults offered Bidens pilosa (cobbler's pegs), Chromolaena<br />
odorata and M. micrantha (all Asteraceae) attacked only the latter. No natural enemies<br />
have been recorded.<br />
The chrysomelid subfamily Chrysomelinae to which this species belongs includes<br />
several successful biological control agents such as the Chrysolina species on St John's<br />
Wort, Hypericum perforaturn angustifolium. If species of Desmogramma are specific to<br />
M. micrantha they may prove to have potential as biological control agents.<br />
Echoma (= Omoplata) marginata and E. quadristillata Coleoptera: Chrysomelidae<br />
Adults and larvae of these chrysomelid beetles feed openly on M. micrantha and<br />
M. cordifolia leaves and cause general defoliation. E. marginata is uncommon, but<br />
E. quadristillata is quite common around Turrialba (Costa Rica). They appear to have a<br />
low reproductive potential (Cock 1982a). In limited-choice tests, E. quadristillata fed on<br />
M. micrantha and M. cordifolia, but not on Bidens pilosa or Sonchus sp. (Asteraceae). In<br />
another experiment, no preference was shown between its two host species, but it would<br />
not feed on another species of Mikania, which was probably M. vitifolia.<br />
A tachinid pupal parasitoid Hyalomyodes triangulifer is known from E. marginata<br />
and a chalcidid pupal parasitoid Brachymeria russelli from E. quadristillata.<br />
Liothrips mikaniae Thysanoptera: Phlaeothripidae<br />
This thrips occurs in Colombia, Costa Rica, Peru, Surinam, Trinidad and Venezuela. The<br />
eggs are mainly laid on the undersurface of the host plant leaves or at the base of leaf<br />
stalks and the larvae feed there in groups. The prepupae and pupae are found among leaf<br />
litter beneath the plant and the adults return to the youngest leaves to feed, mate and<br />
oviposit. L. mikaniae has been found only on M. micrantha growing in full sunshine and<br />
it never occurs on plants in shady situations. This limits its potential effectiveness to sun-<br />
lit stands of the weed. The life cycle (egg to egg-laying adult) takes about 35 days, males<br />
living about 28 days, females about 35 days and laying between 21 and 11 1 eggs (Ooi et<br />
al. 1993). The feeding by larvae and adults on the young leaves produces small to moder-<br />
ate-sized lesions on the undersurface, which dry to form brown scars and these cause<br />
extensive distortion of the leaves as they grow.<br />
Laboratory studies in Trinidad (Cock 1982b) and field observations (Cock 198 1,<br />
1982a,b) show that L. mikaniae is restricted to the genus Mikania and most probably to
4.1 4 Mikania micrantha 131<br />
M. micrantha, although M. cordifolia and M. vitifolia may be fed on to a limited extent in<br />
the laboratory. Additional studies carried out in England by CIBC confirmed its host<br />
specificity before permission was obtained to introduce L. mikaniae to Malaysia. Rearing<br />
methods are described by Cock (1982b) and Ooi et al. (1993).<br />
Physimerus pygmaeus Coleoptera: Chrysomelidae<br />
This halticine chrysomelid is one of a group of five Physimerus species occurring on<br />
M. micrantha in South America.<br />
The larval feeding habits are unknown, but they may attack roots. The adults feed<br />
on young leaves and petioles, causing the dieback of growing tips, and they may be<br />
destructive when in high densities. This species is uncommon in Trinidad, where it is<br />
restricted to shady conditions, whereas in Colombia it also occurs in the open.<br />
Adults of I? pygmaeus have been found feeding on both M. vitifolia and M. hookeri-<br />
ana, in addition to M. micrantha. Field-collected adults fed on Bidens pilosa, but not on<br />
Chromolaena odorata. No natural enemies have been recorded. Further specificity tests<br />
with larvae and adults would be necessary before the potential of this species could be<br />
evaluated. Various halticine beetles, Longitarsus spp., show great promise for the biolog-<br />
ical control of ragwort Senecio jacobaea, Paterson's curse Echium plantagineum and<br />
common heliotrope Heliotropium europaeum.<br />
Pseudoderelomus baridiifomis Coleoptera: Curculionidae<br />
Larvae of this weevil are not known and may be root or stem gall feeders. The adult bur-<br />
rows into the flowers, damaging the petals, stamens and stigma and prevents seed pro-<br />
duction from the flowering head it occupies. When common, levels of 25% damage have<br />
been recorded.<br />
Adults of P. baridiiformis occur mainly in the flowers of M. micrantha, but have<br />
also been recorded from M. trinitaria and M. vitifolia. They occur rarely in the flowers of<br />
Chromolaena odorata and have been recorded once from Neurolaena lobata. No natural<br />
enemies are known.<br />
Although the level of damage caused may be considerable, further studies of life<br />
history and host specificity are required.<br />
Teleonemia sp. or spp. nr prolixa Hemiptera: Tingidae<br />
A taxonomic study of the bug genus Teleonemia (which contains more than 80 species) is<br />
required to enable the determination of correct identity of the one or more species of<br />
brown tingid bugs feeding on Mikania flowering heads. The species is not i? prolixa,<br />
which is highly specific to Lantana camara (Harley and Kassulke 19,759.<br />
The eggs are laid into the flower bracts and the nymphs and adults feed on the<br />
flower heads, but do not appear to cause much damage at low densities. Faeces deposited<br />
on the opening flowers may be sufficient to prevent seed production, particularly when<br />
these serve as a substrate for fungal growth. i? harleyi in Trinidad has a similar life cycle<br />
and feeding habits in Lantana camara flowers (Harley and Kassulke 1975).<br />
Adults and nymphs of Teleonemia were found by Cock (1982a) on a number of<br />
Mikania species (micrantha, vitifolia, trinitaria, hookeriana). Although T. prolixa has<br />
been recorded from Cinchona sp. (Drake and Poor 1938), Lantana camara (Monte 1939)
132 Biological Control of Weeds: Southeast Asian Prospects<br />
and Acacia riparia (Drake and Ruhoff 1965), the records for Cinchona and Acacia<br />
appear to be in error (Harley and Kassulke 1975). A parasite attacks the eggs of<br />
Teleonemia and the lygaeid Xyonysius sp. in M. micrantha flowers. Teleonemia scrupu-<br />
losa has been used in a number of countries to considerable effect to help in the control<br />
of Lantana camara. If the flower-feeding Teleonemia that attack Mikania cause similar<br />
effects through injection of saliva, they may cause more damage than is apparent at first<br />
sight.<br />
MINOR SPECIES<br />
Cock (1982a) and Freitas (1991) list a further 22 species of insects attacking M. micran-<br />
tha in Central and South America (Table 4.14.2). There are 7 species of Hemiptera, 3<br />
Coleoptera, 10 Lepidoptera and 3 Diptera. Five of these are considered at this stage of<br />
knowledge to be promising.<br />
Exema complicata Coleoptera: Chlamisidae<br />
Adults and larvae of this beetle are leaf feeders on Mikania spp., Chromolaena odorata<br />
and perhaps other Asteraceae.<br />
Longitarsus nr amazonus Coleoptera: Chrysomelidae<br />
Adults of this halticine beetle feed on leaves of M. micrantha and larvae probably on<br />
roots. Longitarsus species generally have a very restricted host range.<br />
Neolasioptera sp. Diptera: Cecidomyiidae<br />
The larvae of this fly feed within the flower head and scar the seed shell, but the effect of<br />
this damage on seed viability is unknown. This species is parasitised by a eulophid<br />
Tetrastichus sp.<br />
Sceloenopla sp. Coleoptera: Chrysomelidae<br />
Adults of this hispine beetle feed on leaves and larvae are leafminers on M. micrantha<br />
and M. trinitaria. Horismenus? aeneicollis is recorded as a larval parasitoid. Hispine bee-<br />
tles have proved to be very effective against Lantana camara.<br />
Tegosa claudina Lepidoptera: Nymphalidae<br />
Earlier referred to as Tegosa similis, the larvae of this butterfly are leaf feeders on both<br />
Mikania micrantha and M. cordifolia. Eggs are laid in clusters and larvae are gregarious,<br />
passing through six instars (Freitas 1991).<br />
Attempts at biological control<br />
MALAYSIA<br />
An extensive evaluation of natural enemies attacking M. micrantha was made in<br />
peninsular Malaysia prior to a decision to embark upon a biological control project (Teoh<br />
et al. 1985). Of the insects collected from or reared on the host plant, 2506 were classi-<br />
fied arid separated into commonly found and minor natural enemies (Table 4.14.3).<br />
Although numerous insects were found to attack M. micrantha, not only was the<br />
extent, of damage insignificant, but most of the abundant species were known pests of<br />
economic crops. For example, Halticus minutus and Lamprosema diemenalis are major
4.14 Mikania micrantha 133<br />
pests of leguminous cover crops, Homoeocerus serrifer attacks rice and Helopeltis spp.<br />
are serious pests of cocoa. None of the natural enemies recorded in Tropical and South<br />
America were found in the survey. A number of fungi were also found, of which<br />
Colletotrichium gloeosporioides was the most important, comprising 84% of the sam-<br />
ples. Other fungi included Colletotrichium spp., a non-sporulating brown fungus,<br />
Rhizoctonia spp., Curvularia spp. and Pestalotia spp. It was concluded that a strong case<br />
existed for the introduction of effective natural enemies.<br />
Table 4.14.3 Insects attacking Mikania micrantha in peninsular Malaysia (Teoh et al.<br />
1985).<br />
Species Effects<br />
MAJOR SPECIES<br />
Orthoptera<br />
ACRlDlDAE<br />
Acrida turrita<br />
Hemiptera<br />
APHlDlDAE<br />
Aphis spiraecola<br />
CERCOPIDAE<br />
Clovia conifer<br />
CICADELLIDAE<br />
Bothrogonia ferrugenea<br />
COREIDAE<br />
Homoeocerus serrifer<br />
Riptortus linearis<br />
MEMBRACIDAE<br />
Centrotypus flexuosus<br />
MlRlDAE<br />
Halticus minutus<br />
Helopeltis spp.<br />
Lepidoptera<br />
PY RALl DAE<br />
MINOR SPECIES<br />
Orthoptera<br />
ACRlDlDAE<br />
Catantops humilis<br />
Hemiptera<br />
APHlDlDAE<br />
Aphis gossypii<br />
CICADELLIDAE<br />
Nephotettix spp.<br />
Thysanoptera<br />
THRlPlDAE<br />
Isothrips spp.<br />
Microcephalothrips spp.<br />
holes in leaves<br />
wrinkled leaves<br />
yellow spots on leaves and stems<br />
brown spots on stems<br />
brown spots on leaves<br />
brown spots on leaves<br />
necrosis on stems<br />
necrotic lesions on leaves<br />
necrotic lesions on leaves<br />
(continued on next page)
134 Biological Control of Weeds: Southeast Asian Prospects<br />
Table 4.14.3 (continued)<br />
Species Effects<br />
Parthenothrips spp.<br />
Thrips hawaiiensis<br />
Thrips tabaci<br />
Coleoptera<br />
CHRYSOMELIDAE<br />
Dactylispa bipartista<br />
COCCINELLIDAE<br />
Coelophora bissellata<br />
Epilachnu indica<br />
Diptera<br />
TEPHRlTlDAE<br />
Sphaeniscus atilus<br />
Lepidoptera<br />
AMATIDAE<br />
Amata huebneri<br />
PYRALIDAE<br />
Hellula undalis<br />
Liothrips mikaniae was introduced to Malaysia from Trinidad via England in 1989<br />
for additional host specificity trials. Difficulties were experienced initially in mass rearing,<br />
due to unsuitable environmental conditions (lighting, temperature, aeration) low<br />
plant nutritional quality and predators (spiders, ants, and particularly a predatory thrips,<br />
Xyloplothrips sp. which destroyed 90% of the culture). Also, there were differences in<br />
the M. micrantha plants used. Some were hairless and others hairy. Larvae hatching from<br />
eggs along stems of the latter found difficulty in moving among the trichomes and many<br />
failed to reach the nearest leaf. Nevertheless, in Trinidad, L. mikaniae was found breeding<br />
on both plant types. After passing tests with 18 Malaysian crop species, 13 161 adult<br />
thrips were released in 25 batches of 99 to 1400 at 5 different sites from April 1990 to<br />
June 1991, but no establishment resulted (Table 4.14.4). Further thrips were imported in<br />
January 1992 to extend the genetic base of the rearing colony. Two releases were made<br />
into a fenced site, 18 000 adults, together with their rearing plants in pots, in May 1992<br />
and 1500 adults about a month later. The pots were watered daily for two months.<br />
However, L. mikaniae gradually disappeared and, after eight months, none could be<br />
found. An ant that made its nest amongst Mikania leaves was observed to feed voraciously<br />
on adult, larval and pupal stages of L. mikaniae, but showed little interest in eggs,<br />
whereas a predatory Haplothrips sp. preferred eggs. About 20 other potential predators<br />
were evaluated, but were not implicated (Liau et al. 1991, 1993, Norman et al. 1992,<br />
Teoh et al. 1985, Ooi 1993, Ooi et al. 1993).<br />
SOLOMON ISLANDS<br />
A consignment of L. mikaniae was sent from Malaysia to the Solomon Is and released in<br />
the field in 1988, but the site was flooded shortly afterwards. Further releases were made,<br />
but no establishment has occurred (M. Vagalo pers. comm.). It was suggested that there
4.14 Mikania micrantha 135<br />
may be differences in hospitability to L. mikaniae of the host plant between the<br />
Caribbean and Solomon Is.<br />
PAPUA NEW GUINEA<br />
Part of a consignment of L. mikaniae sent to the Solomon Is in 1989 was taken to Papua<br />
New Guinea, but there is no information on its fate (Williams et al. 1990), although it is<br />
believed to have died in quarantine (F. Dori pers. comm. 1993).<br />
SRI LANKA AND ASSAM<br />
In Sri Lanka and Assam it has been found that the plant parasite Cuscuta chinensis will<br />
suppress Mikania and prevent it spreading into tea plantations, although C. chinensis is<br />
not sufficiently selective to be used in the plantations themselves (Parker 1972). On<br />
Espiritu Santo (Vanuatu) a related species Cuscuta campestris is reported to suppress M.<br />
micrantha (M.J.W. Cock pers. comm.).<br />
Table 4.14.4 Liberations for the biological control of Mikania micrantha.<br />
Species Where From When Result References<br />
Thysanoptera<br />
PHLAEOTHRIPIDAE<br />
Lioihrips mikaniae Malaysia Trinidad via UK 1991 - Liau et al. 1991,<br />
1993, Norman et<br />
al. 1992<br />
Solomon Is Trinidad via UK 1988 - M. Vagalo pers.<br />
comm., Williams et<br />
al. 1990<br />
Comment<br />
Mikania micrantha is an introduced weed of widespread importance in plantation crops<br />
throughout Southeast Asia. It is not a significant weed in its native range in tropical<br />
Central and South America, where it is attacked by a wide range of arthropod natural<br />
enemies. Several appear to be not only damaging to the weed, but also highly specific. It<br />
must be concluded that M. micrantha is a highly appropriate target for an attempt at bio-<br />
logical control.<br />
The failure of Liothrips mikaniae to become established in Malaysia is disappoint-<br />
ing. The most probable explanation, partly supported by observations, is that heavy pre-<br />
dation was the cause. Although the same guild of predators may be widespread on conti-<br />
nental Southeast Asia, the same may not apply to island nations, particularly those in the<br />
southern Pacific, which have a far less diverse fauna and releases there might lead to suc-<br />
cessful establishment.
136 Biological Control of Weeds: Southeast Asian Prospects<br />
Mimosa in visa<br />
(after Holm eta/. 1977)
Map 4.15 Mimosa invisa<br />
4.15 Mimosa invisa<br />
Mimosa in visa<br />
Creeping sensitive plant, Mimosa invisa, is native to Tropical America, where it is not<br />
regarded as a weed.<br />
At least 70 species of insects attack it in Brazil and additional species elsewhere.<br />
Detailed studies have been made on two Hemiptera (Heteropsylla spinulosa and<br />
Scamurius sp.) and a moth (Psigida walkeri). H. spinulosa has caused extensive damage<br />
to M. invisa following its establishment in Australia and promising early results in Fiji,<br />
Papua New Guinea and Pohnpei, but disappointing results in Western Samoa. Scamurius<br />
sp. has failed to become established and P. walkeri is still under investigation.<br />
The prospects for biological control of M. invisa appear to be good, although<br />
additional natural enemies may have to be considered.
138 Biological Control of Weeds: Southeast Asian Prospects<br />
4.15 Mimosa invisa Martius ex Colla<br />
Mi mosaceae<br />
creeping sensitive plant; banla saet (Cambodia), borang, pis koetjing, rembete<br />
(Indonesia), duri semalu (Malaysia), makahiyang lalaki (Philippines), maiyaraap<br />
thao (Thailand), co trinh nu moc (Vietnam)<br />
Rating<br />
+++ Phil<br />
18 ++ Myan, Thai, Laos, Viet, Msia, Sing, Indo<br />
+ Camb<br />
Origin<br />
Tropical America. In Brazil southwards from Bahia to Paranh and westwards to Paraguay<br />
and tropical northeast Argentina; also lowlands of Central America from Veracruz<br />
(Mexico) southeastwards to Panama and adjacent Colombia.<br />
Distribution<br />
In addition to the above there are, in the Americas, scattered records from Brazilian<br />
Amazonia, the Guianas, Jamaica, Hispaniola and Cuba. M. invisa is widely distributed in<br />
Southeast Asia and the Pacific, also in Queensland, India, Sri Lanka, Taiwan and Nigeria.<br />
It was first recorded in Java in 1900 (Soerjani et al. 1987).<br />
Characteristics<br />
Mimosa invisa is a fast growing, abundantly thorny, biennial or perennial shrub with<br />
angular branching stems that become woody with age. The leaves are alternate, bipinnate<br />
and compound. The pink to purple globular flowers are borne on a short prickly stalk<br />
arising from a leaf axil. The seed pods are covered with stiff bristles and separate at<br />
transverse grooves into two to four, single-seeded segments.<br />
The genus Mimosa does not occur naturally in Southeast Asia or Australia. M.<br />
invisa is one of three weedy species of Mimosa in this region, all of which are treated in<br />
this volume. They may be distinguished (i) by the number of pinnae in the leaves: M.<br />
invisa 4 to 9 pairs; M. pigra 6 to 16 pairs; and M. pudica 1 to 2 pairs and (ii) the size of<br />
the pods: M. invisa 4 seeds per pod, M. pigra 12 to 24 seeds. In addition, M. invisa stems<br />
have a dense covering of small prickles, whereas M. pigra stems have a sparse covering<br />
of large prickles (Lonsdale 1992).<br />
M. invisa folds its pinnate leaves when touched, but is not as sensitive as some other<br />
species, such as M. pudica. The leaves fold at nightfall.<br />
unlike the situation in the more tropical regions, such as the Philippines where<br />
M. invisa flowers all year round, in central and southern Brazil it only flowers from the<br />
end of January to mid April. Seeds mature from February to the end of May and plants<br />
then senesce, losing most of their leaves, although a few green leaves remain at the stem
4.15 Mimosa invisa 139<br />
base. For two to five months green plants are difficult to find. Senescence is not due to<br />
water shortage as well-watered plants in the laboratory also senesce. However, germina-<br />
tion occurs when moisture is available, so young plants may appear after showers of rain<br />
(Garcia 1982b).<br />
Importance<br />
M. invisa scrambles vigorously over other plants, forming dense tangled thickets up to 2<br />
m high. It is a nitrogen-fixer and its extremely rapid growth smothers useful plants and<br />
other weeds. Its sharp, recurved thorns make stock reluctant to graze on it and difficult<br />
for them to penetrate the stands. Crops infested with M. invisa are difficult to harvest<br />
because the thorns puncture and lacerate the hands of the workers. It is common along<br />
roadsides and in moist waste places. It causes major problems in coconut, tea and rubber<br />
plantations, sugarcane and pineapple fields, crop lands and pastures. It is not a problem<br />
in the Americas, western Asia, East Africa or Europe, but is a serious weed in Southeast<br />
Asia and the Pacific. A spineless variety, M. invisa inermis, has been suggested as a trop-<br />
ical pasture legume, but its tendency to revert to the thorny type and its potential toxicity<br />
has discouraged its use (Waterhouse and Norris 1987).<br />
In the Americas, M. invisa is most common in the Paranh basin in the State of San<br />
Paulo (Brazil), but even there pure stands are not common and it does not appear to<br />
invade nearby crops. In forest regions it occurs as thickets among grasses along roads,<br />
river banks and in waste places; it occurs more commonly on the fringe of cities (Garcia<br />
1982b).<br />
Natural enemies<br />
Information up to 1986 was summarised by Waterhouse and Norris (1 987).<br />
M. invisa is seldom troublesome in Brazil or Argentina and some 70 species of<br />
insects and two fungi are known to attack it in Brazil (Table 4.15.1). Additional insects are<br />
known in the Americas, but they have not yet been studied. A pathogenic fungus<br />
Corynespora cassiicola kills M. invisa in Australia (Haseler 1984), heavy infestations of<br />
scale insects attack it in Fiji (Mune and Parham 1967), a non-specific lymantriid larva<br />
feeds on young leaves and flowers in Thailand (Napompeth 1982) and a pierid butterfly<br />
Eurema sp. breeds on it in Papua New Guinea and New Britain (T.L. Fenner pers. comm.).<br />
More than half of the insects attacking M. invisa in Brazil have not yet been identi-<br />
fied. Indeed it is probable that most of these are undescribed species and, if so, it follows<br />
that there is no published information about them. Where possible, identification, even to<br />
a genus, may be valuable. For example, species of the genus Heteropsylla are restricted<br />
to legumes, with known hosts only in the Mimosaceae or Caesalpiniaceae. Of the 35<br />
described species with recorded hosts, 31 are specific to a single host (Hodkinson and<br />
White 1981, Muddiman et al. 1992).<br />
Only a few of the 70 insect species attacking M. invisa are known as agricultural<br />
pests (Table 4.15.1). From the remainder, preliminary observations on a subgroup of<br />
about 10 species led to detailed studies on three, Heteropsylla spinulosa, Scamurius sp.<br />
and Psigida (= Psylopigida) walkeri.
140 Biological Control of Weeds: Southeast Asian Prospects<br />
Attempts at biological control<br />
AUSTRALIA<br />
Large numbers of Heteropsylla spinulosa were released in coastal Queensland, com-<br />
mencing in 1988 (Table 4.15.2). This involved 33 field sites, averaging thousands of<br />
insects per release (M. Ablin pers. comm. 1990). Within two years, the psyllid had dis-<br />
persed widely into all infestations of M. invisa in pastures. Dense clumps of the weed<br />
were reduced to small masses of bare stems with stunted growing tips, leading to other<br />
vegetation reestablishing itself. Seed production from severely affected plants was<br />
reduced by 85 to 100% (Ablin 1992, Anon 1988). It did not attack M. pudica plants near-<br />
by. A more recent assessment, using insecticide-produced exclusion, found that H. spinu-<br />
losa reduced seed production on average by SO%, growing tip elongation by 77% and the<br />
growth rate of tips by 50% (Ablin 1993a). Although M. invisa may still produce clusters<br />
of seed pods when damage is high, the pods contain very few viable seeds. Mature plants<br />
support low populations of H. spinulosa during the dry season from July to November.<br />
Thereafter, psyllid abundance increases with the onset of summer rains, with peak num-<br />
bers in April or May (M. Ablin pers. comm. 1993).<br />
Scamurius sp. was liberated in Queensland from 1987 to 1990 and proceeded to kill<br />
the tips of many shoots (Anon 1988). However it did not become established. It was also<br />
released against Mimosa pigra in the Northern Territory where it survived for several<br />
months, but fecundity was very low and the colony died out (M. Ablin pers. comm.<br />
1993).<br />
FIJI<br />
H. spinulosa was brought in from both Western Samoa and Australia in 1993 and, after a<br />
generation in quarantine, was liberated in Nadi in June. Six weeks later all stages were<br />
seen in the field. A mealy bug and Tetranychus sp. mites are occasionally found on<br />
M. invisa in the field (S.N. La1 pers. comm. 1993).<br />
PAPUA NEW GUINEA<br />
Heteropsylla spinulosa from Australia was reared through one generation in quarantine<br />
in Port Moresby and released early in 1993 in the Ramu Valley near Lae. Within a few<br />
months it had severely damaged M. invisa and killed many plants (F. Dori pers. comm.<br />
1993).<br />
POHNPEI<br />
Ten months after release at Palikir, Pohnpei H. spinulosa became abundant on M. invisa<br />
and subsequently killed many plants. Many psyllids were transferred to M. invisa in other<br />
areas (N.M. Esguena pers. comm. 1993).<br />
WESTERN SAMOA<br />
A total of 47000 nymphs and adults of Heteropsylla spinulosa from Australia were liber-<br />
ated in Western Samoa in 1988 and 1989 and, a year later, the psyllid was reported at<br />
some sampling sites to have reduced seed production, although not the area infested<br />
(Willson and Ablin 1991).<br />
Scamurius sp. was also liberated in Western Samoa in 1989 and was seen in the<br />
field after more than one generation, but not in more recent times. There have been no
4.15 Mimosa invisa 141<br />
reports of its effects. M. invisa continues to be a serious weed, with more than 85% of<br />
villages on the main island of Upolu being infested (Willson and Garcia 1992).<br />
Important Natural Enemies<br />
INSECTS<br />
Heteropsylla spinulosa Hemiptera: Psyllidae<br />
The average development period of this small (2.5 mm long) pale green, Brazilian psyllid<br />
is about 28 days. High populations cause severe stunting and distortion of the leaves and<br />
growing tips; flowering is reduced or even prevented. A sticky honeydew is produced<br />
which encourages a dense growth of sooty moulds. Females attach eggs by means of a<br />
pedicel inserted into the plant tissue between overlapping leaflets. Young nymphs live<br />
hidden between the leaflets, whereas adults feed on leaflets and shoots.<br />
In Brazil reduviid bugs pierce nymphs with their proboscis and withdraw them from<br />
their shelters, whereas larvae and adults of the predatory coccinellid Eriopis connexa are<br />
only able to capture exposed nymphs. Nymphs are also attacked by an encyrtid wasp<br />
Psyllaephagus yaseeni (Willson and Garcia 1992). The predatory vespid wasp<br />
Protonectarina sylveiriae attacks nymphs and an unidentified wasp causes up to 13%<br />
parasitisation (Garcia 1985).<br />
In extensive host specificity tests H. spinulosa adults and nymphs were unable to<br />
live on any plant other than M. invisa and its spineless variety M. invisa inermis. In the<br />
field it did not attack M. pudica, even when large infestations of M. invisa were<br />
destroyed and M. pudica was common nearby (M. Ablin pers. comm. 1993). Eggs were<br />
laid on 18 other plant species, but only under glasshouse conditions (Garcia 1985, Wild<br />
1987, Willson 1987, Willson and Garcia 1992) and Heteropsylla spinulosa was judged<br />
safe to liberate in Australia and four other countries (Table 4.15.2)<br />
Psygida walkeri Lepidoptera: Cercophanidae<br />
This moth is widespread in Brazil and Colombia. Females have a wing span of up to 5<br />
cm. When fully grown its greenish, spiny larvae may reach a length of 5 cm. They feed<br />
voraciously on leaves, flower buds, tender seed pods and on the top 30 cm of tender<br />
stems and branches, preventing both flowering and seed production. The life cycle takes<br />
about 2 months and there are 3 generations a year. There is a pupal diapause of up to 4<br />
months in Brazil<br />
Larvae of P. walkeri have been found in the field on M. invisa, M. rixosa, M. vel-<br />
loziana and once on M. somnians. They have not been found in the field in Brazil on<br />
other leguminous plants near M. invisa plants bearing larvae, nor on any plants of eco-<br />
nomic importance (Garcia 1983). However, under artificial conditions larvae can be<br />
reared on black wattle Acacia mearnsii and may also feed on Mimosa pudica and<br />
Leucaena leucocephala. Although adults will oviposit on A. mearnsii, no attack has been<br />
observed in the field (Haseler 1984). Further host specificity testing is in progress in<br />
Australia.
142 Biological Control of Weeds: Southeast Asian Prospects<br />
Scamurius sp. Hemiptera: Coreidae<br />
Both nymphs and adults of this large (up to 2.2 cm) coreid bug feed on the shoots, caus-<br />
ing them to collapse, thereby inhibiting vegetative growth and flowering. First instar<br />
nymphs moult whether fed or not and, after five instars, mature in about seven weeks.<br />
There are about four generations a year, from early summer to autumn, and adults may<br />
live for six months or more.<br />
Adults were found to probe many species of plants, but to feed only on species of<br />
Mimosa. Nymphs were able to develop on M. invisa and on two other weedy species,<br />
M. pigra and M. pudica, but not on other plants (Garcia 1984, Wild 1986, 1987). This<br />
species was approved for release in Australia and Western Samoa (Table 4.15.2).<br />
FUNGUS<br />
Corynespora cassiicola<br />
This stem spot fungus is very common in hot humid weather in north Queensland, Papua<br />
New Guinea (Keravat, Rabaul) and Western Samoa. It can be very damaging to M. invisa<br />
if environmental conditions are suitable (Willson and Ablin 1991). The strain involved<br />
appears to be specific to M. invisa, although fungi with the same specific name are<br />
reported from cowpea, papaya and tomato. If suitable environmental conditions persist in<br />
the field M. invisa plants shed their leaflets and stems die back as lesions cover the plant<br />
(M. Ablin pers. comm. 1993).<br />
Comments<br />
The use, in the future, of H. spinulosa from Brazil against M. invisa is complicated by<br />
several introductions of natural enemies that have already been made by Southeast Asian<br />
countries (eg. Thailand, Philippines). These introductions were of two parasitic wasps<br />
(Tamarixia leucaenae and Psyllaephagus yaseeni) native to the Caribbean and Central<br />
America (Noyes 1990) and one or more predatory coccinellids against a pest psyllid<br />
Heteropsylla cubana which appeared from the Americas in the 1980s. This pest can<br />
cause severe damage to Leucaena leucocephala which is widely planted for firewood and<br />
as fodder. Tests showed that these natural enemies of H. cubana would also attack<br />
H. spinulosa (Baker 1990). As a result, several countries where M. invisa is a serious pest<br />
(Australia and most Pacific islands, but not New Caledonia) have deferred introducing<br />
natural enemies of H. cubana.<br />
Several interesting points, relevant to any investigation for natural enemies in<br />
South America, emerge from the M. invisa project there:<br />
Before the studies of C.A. Garcia in the early 1980s almost nothing was known about<br />
the insects attacking it, yet within a year 57 insects were listed from Brazil (Garcia<br />
1982a,b) and within two years a further 10. No records are available from elsewhere<br />
in the Americas, except of a Heteropsylla sp. from Colombia (Garcia 1983).<br />
It has not been possible for taxonomists to assign a specific name (and sometimes not<br />
even a generic name) to the majority of insects collected. Some were not previously<br />
represented in any museum collections and many are almost certainly undescribed<br />
species.
4.1 5 Mimosa invisa 143<br />
Only one (Scamurius sp.) of the three insects eventually selected for detailed study<br />
was recognised in the first survey which yielded 57 species. It is clear that follow up<br />
surveys are essential.<br />
No study has yet been made of the insects attacking M. invisa over a very large area<br />
of its presumed native range. From brief observations in Colombia Garcia (1 983)<br />
commented that larvae of Lepidoptera were 'quite similar to those collected off M.<br />
invisa in Brazil. Coleoptera in general look different'.<br />
Should existing biological control be considered inadequate, further detailed studies<br />
covering the entire native range of M. invisa might well reveal additional, adequately<br />
specific insects attacking it.<br />
Examination of the host specificity of more of the insects recorded from Brazil (Table<br />
4.15.1) might also reveal further adequately specific insects.<br />
H. spinulosa does not thrive under either very wet or very dry conditions. Its popula-<br />
tions depend upon the availability of green foliage and, in the dry season, are found<br />
on pockets of green foliage. A flush of growth after rain leads to a build up in popula-<br />
tions to a level that severe damage may be caused, sufficient to kill many M. invisa<br />
plants.<br />
Table 4.15.1 Natural enemies of Mimosa invisa in Brazil (from Garcia 1982a,b, 1983<br />
and his unpublished monthly reports).<br />
Species Comment<br />
INSECTS<br />
Orthoptera<br />
TETTlGONl IDAE<br />
sp. 1<br />
sp. 2<br />
attacks flowers<br />
attacks leaves<br />
Hemiptera<br />
CERCOPIDAE<br />
Tomaspis (= Zulia) enteriana<br />
SP . belongs to Gyopinae<br />
CICADELLIDAE<br />
sp. I common on Mimosa invisa and M. pigra and<br />
colonises Acacia mearnsii and M. scabrella during<br />
the dry season<br />
sp. 2<br />
COREIDAE<br />
Scamurius sp. 1 see text<br />
Scamurius sp. 2<br />
MlRlDAE<br />
Horciacinus signoreti (= H. argentinus)<br />
Taylorilygus pallidulus<br />
MEMBRACIDAE<br />
Ceresa ustulata<br />
an agricultural pest<br />
Enchenopa gracilis<br />
Micrutalis sp.<br />
(continued on next page)
144 Biological Control of Weeds: Southeast Asian Prospects<br />
Table 4.15.1 (continued)<br />
Species Comment<br />
PENTATOMI DAE<br />
Acrosternum herbidum<br />
Dichelops furcatus an agricultural pest<br />
Edessa meditabunda an agricultural pest<br />
Euschistus tristigmus cribarius<br />
Euschistus luridus<br />
Piezodorus guildinii an agricultural pest<br />
PSYLLIDAE<br />
Heteropsylla spinulosa see text<br />
Heteropsylla sp. from Colombia<br />
THYREOCORIDAE<br />
Gyrocnemis sp.<br />
Coleoptera<br />
CHRYSOMELIDAE<br />
Colaspis sp.<br />
Cryptocephalus viridiaeneus<br />
?Hilax sp.<br />
Lactica sp.<br />
Lexiphanes ?semicyaneus<br />
Lexiphanes sp.<br />
Metaxyonycha pallidula<br />
Nodonota sp.<br />
Pachybrachys sp.<br />
System s-liftera<br />
Temmodachrys sp. nr aphodoides<br />
CURCULIONIDAE<br />
Asynonychus godmani<br />
(= Pantomorus cervinus)<br />
Chalcodermus sp.<br />
Chalcodermus sp. nr segnis<br />
Hypanthus sp.<br />
Promecops sp. 1<br />
Promecops sp. 2<br />
Sibinia aspersa<br />
Sibinia ?subulirostris<br />
Sibinia sp.<br />
Lepidoptera<br />
AMATIDAE<br />
SP.<br />
CERCOPHANIDAE<br />
Psigida walkeri<br />
GEOMETRIDAE<br />
sp. 1<br />
sp. 2<br />
most Chrysomelidae were present ib low numbers.<br />
adults eat M. invisa leaves<br />
excellent defoliator; also attacks M. pigra<br />
adults eat leaves and show strong preference for<br />
M. invisa, but will also attack M. pudica, Calliandra<br />
selloi and Acacia mearnsii<br />
larvae bore stems, brown adults eat leaves<br />
black adults bore green seeds, but did not attack pods<br />
of 7 other legumes<br />
adults eat ovaries, larvae the seeds; recorded from<br />
Mimosa albida and M. quadrivalis, but would not<br />
attack six other legumes including Leucaena<br />
leucocephala<br />
seminicola group, larvae eat seeds, adults the leaves<br />
and ovaries and are also found in Mimosa rixosa<br />
red hairy larva<br />
see text<br />
debris-covered larva<br />
twisted larva<br />
(continued on next page)
FUNGI<br />
Species Comment<br />
sp. 3<br />
sp. 4<br />
sp. 5<br />
HESPERIIDAE<br />
Cogia (= Caicella) calchas<br />
LYCAENIDAE<br />
Hemiargus hanno<br />
Tmolus (= Thecla) azia<br />
NOCTUIDAE<br />
sp. 1<br />
sp. 2<br />
sp. 3<br />
sp. 4<br />
PlERlDAE<br />
Eurema tenella<br />
TORTRICIDAE<br />
sp. 1<br />
sp. 2<br />
sp. 3<br />
sp. 4<br />
FAMILY UNKNOWN<br />
Cercospora canescens<br />
Fusarium sp.<br />
Uredo mimosae-invisae<br />
reddish green larva<br />
common slim larva<br />
thick twig larva<br />
Table 4.15.2 Liberations for biological control of M. invisa.<br />
4.1 5 Mimosa invisa 145<br />
occurs in Mexico and Argentina; eggs laid also on<br />
M. pudica, M. scabrella, Indigofera anil, Skranquia<br />
sp., larvae are heavily parasitised<br />
larvae eat leaves, flowers and seed pods; also<br />
M. pudica flowers<br />
larvae eat leaves and flowers; also M. pudica and<br />
groundnut flowers<br />
velvet black larva; also on M. pudica, M. scabrella,<br />
Acacia mearnsii, Calliandra selloi, Leucaena<br />
leucocephala<br />
reddish larva<br />
green larva<br />
small green larva<br />
occurs in Brazil, Argentina, Paraguay, Bolivia; larvae<br />
defoliate M. invisa; also eat M. pudica, M. scabrella,<br />
Acacia mearnsii and (reluctantly) Leucaena<br />
leucocephala<br />
flowerltwig roller<br />
pod eater<br />
larvae eat leaves<br />
larvae roll flowers<br />
4 species, two bore in the stem tips, 1 eats pods<br />
and I the flowers<br />
from Venezuela (H.C. Evans pers. comm. 1992)<br />
Species From Liberated When Result References<br />
Heteropsylla spinulosa Brazil Queensland 1988 + Ablin 1992, Anon 1988<br />
Fiji 1993 ? S.N. Lal pers. comm. 1993<br />
Papua New Guinea 1993 + Ablin 1993b,<br />
F. Dori pers. comm. 1993<br />
Pohnpei 1992 + N.M. Esguerra pers. comm.<br />
1993<br />
Western Samoa 1988 + Willson & Garcia 1992<br />
Scamurius sp. Brazil Queensland 1987 - Anon 1988<br />
Western Samoa 1988 ?
146 Biological Control of Weeds: Southeast Asian Prospects<br />
Mimosa pigra<br />
(after CSIRO. 1992)
Map 4.16 Mimosa pigra<br />
4.16 Mimosa pigra<br />
Mimosa pigra originated in the area extending from Mexico to Amazonia and<br />
Venezuela. Four beetles and two moths have been established on M. pigra in Australia<br />
in the past 10 years. Two stem-boring moths Neurostrota gunniella and Carmenta<br />
mimosa are having a significant effect on the vigour of the weed. N. gunniella has<br />
spread widely, infests most stems and is reducing seed production. All except N. gun-<br />
niella have been liberated in Thailand. N. gunniella was not liberated because it can<br />
attack the aquatic vegetable Neptunia oleracea. However the two seed-feeding bruchids<br />
are destroying up to 20% of the seed in Thailand<br />
A specific, highly pathogenic fungus, Phloeospora mimosae-pigrae, has been<br />
approved for release in Australia and six insects and a rust fungus are under study.<br />
There are grounds for confidence that a group of natural enemies will become avail-<br />
able that, acting together, will cause significant damage to M. pigra.
148 Biological Control of Weeds: Southeast Asian Prospects<br />
4.16 Mimosa pigra L.<br />
Mimosaceae<br />
giant sensitive plant; mai yah raap yak, maiyarap ton, chi yop luang (Thailand);<br />
kembang gajah, semalu gajah (Malaysia); trinh nu nhon (Vietnam); putri malu<br />
(Indonesia)<br />
Rating<br />
+++ Thai<br />
15 ++ Myan, Laos, Msia, Sing, Indo<br />
+ Camb, Viet<br />
Origin<br />
Mexico, southern Venezuela, or central Amazon basin.<br />
Distribution<br />
Throughout the tropics and still spreading. It was an early invader of tropical Africa and<br />
is spreading aggressively in northern Australia and Southeast Asia. It is not present in the<br />
Philippines or the oceanic Pacific. Details of its distribution and time of recognition in<br />
various countries are given in Lonsdale (1992).<br />
Characteristics<br />
M. pigra is a perennial leguminous shrub, growing up to 6 m high on a wide range of<br />
soils, and found in moist open sites with a rainfall between 750 and 2250 mm in the trop-<br />
ics. Its leaves are bipinnate and sensitive to the touch, through movements of the petiole<br />
and pinnules. Petioles bear a slender prickle at the junction of each of the 6 to 16 pairs of<br />
pinnae and sometimes have stouter prickles between each pair. The stems bear<br />
broad-based, sharp thorns up to 7 mm long. Mature plants have many branches growing<br />
from the base, with a skirt of adventitious roots forming in seasonally inundated sites.<br />
They have a large central taproot which penetrates 1 to 2 m deep and a lateral root system<br />
that extends up to 3.5 m from the stem at a depth of about 5 cm. The flowers are mauve<br />
to pink, massed in globular heads lcm in diameter, with each head containing about 100<br />
flowers. Seed pods are produced in clusters of about 7, are densely bristly, 3 to 8 cm long<br />
and break transversely into segments each containing a seed. The bristles facilitate float-<br />
ing and thus rapid spread of the weed along river systems. In regions with pronounced<br />
wet and dry seasons, the former is the main period of growth, with flowering mainly<br />
from mid to late wet season. Development from flower bud to ripe seed takes about 5<br />
weeks (Lonsdale 1992). Average seed production is about 9000 seeds per plant, but up to<br />
220000 has been recorded. Although most seeds that lodge on or near the soil surface<br />
probably germinate within two years many seeds deeper in the soil lie dormant for long<br />
periods (at least 23 years).<br />
Previously, two varieties were recognised var. pigra and var. berlandieri, of which<br />
only pigra has spread around the world. Variety berlandieri has recently been renamed<br />
Mimosa asperata (Barneby 1989, Lonsdale 1992).
Importance<br />
4.16 Mimosa pigra 149<br />
In tropical America M. pigra usually occurs as small clumps of multi-stemmed plants<br />
growing in seasonally flooded habitats. However, in many countries to which it has been<br />
introduced, M. pigra is a serious weed of wetlands. Dense, prickly thickets compete with<br />
pastures, prevent access to water and hinder mustering. The thickets exclude native vege-<br />
tation and so alter the environment that many native plants and animals are eliminated or<br />
seriously affected (Lonsdale 1992). The weed leads to sediment accumulation in irriga-<br />
tion systems and reservoirs and, as the seed segments float, many end up in fallow rice<br />
paddies where they germinate rapidly. River sand containing seeds helps to establish new<br />
infestations when transported to building sites, road constructions etc.<br />
Cattle and horses occasionally browse on young plants and some wild animals find<br />
it acceptable, particularly as a dry-season browse but, in general, it seems to be of low<br />
palatability. The leaves contain low levels of the toxic amino acid mimosine. M. pigra<br />
was introduced to Thailand in 1945 as a green mulch crop and for erosion control in rice<br />
paddy irrigation channels, but it was soon found that the problems associated with it far<br />
outweighed any advantages (Wara-Aswapati 1983). However it is used still as a source of<br />
firewood and bean poles, although it is now regarded as a very serious weed.<br />
Natural enemies<br />
Surveys for natural enemies have been made in Brazil, Mexico, Venezuela (Harley et al.<br />
1983), Honduras (Habeck and Passoa 1982), Costa Rica (Forno 1992) and most recently<br />
in Belize and Cuba (I.W. Forno pers. comm.). In Honduras more than 60 species were<br />
listed (2 Orthoptera, 27 Hemiptera, 1 Diptera, 15 Coleoptera and 15 Lepidoptera).<br />
Although a full list of insects attacking M. pigra in its native range has yet to be pub-<br />
lished, a diverse group of over 200 is known to occur (Forno et al. 1991b). Only 12<br />
species are considered to be pests of agriculture and at least 45 have habits that are likely<br />
to lead to restricted host specificity, such as gall forming, leaf mining or stem boring. It is<br />
suggested that some 10% may be adequately host specific. These are likely to attack dif-<br />
ferent parts of the plant causing complementary damage, so that the prospects for biolog-<br />
ical control appear good (Forno et al. 1989b, Wilson et al. 1990).<br />
Six of the tropical American species of insects have been liberated (Table 4.16.1)<br />
five more were eventually not released after tests showed (or suggested) that they are<br />
insufficiently host specific (Table 4.16.2) and a further six are currently being examined<br />
in Australia or Mexico for host specificity (Table 4.16.3). However, the list of potential<br />
insects for consideration is far from exhausted. Host testing of agents for M. pigra is dis-<br />
cussed by Forno and Harley (1 992).<br />
Two fungal pathogens of M. pigra (Diabole cubensis and Phloeospora mimosae-<br />
pigrae) cause considerable damage in Mexico in spite of being extensively attacked by<br />
hypeiparasitic fungi. Without these, the pathogens should prove even more damaging.<br />
Other, less damaging fungi include Colletotrichium gloeosporioides, Pestalopsis sp.,<br />
Phomopsis sp. and Oidium sp. (Evans 1988, 1990, Evans and Seier 1991, Evans et al.<br />
1993).
150 Biological Control of Weeds: Southeast Asian Prospects<br />
Table 4.16.1 Releases for the biological control of Mimosapigra.<br />
Species Part attacked Liberated Result References<br />
Coleoptera<br />
APlONlDAE<br />
Coelocephalapion<br />
aculeatum<br />
BRUCHIDAE<br />
Acanthoscelides<br />
puniceus<br />
Acanthoscelides<br />
quadridentatus<br />
CHRYSOMELIDAE<br />
Chlamisus<br />
mimosae*<br />
Lepidoptera<br />
GRAClLLARllDAE<br />
Neurostrota<br />
gunniella<br />
flower buds<br />
seeds<br />
seeds<br />
pinnae &<br />
stems<br />
pinnules &<br />
stems<br />
Australia 1992<br />
Thailand 1991<br />
Australia 1983<br />
Thailand 1984<br />
Vietnam 1987<br />
Australia 1983<br />
Thailand 1984<br />
Vietnam 1987<br />
Australia 1985<br />
Thailand 1986<br />
Vietnam 1990<br />
SESllDAE<br />
Carmenta mimosa stem<br />
Australia 1989<br />
Thailand 199 1<br />
* Introduced from Brazil, the remaining 5 insects from Mexico.<br />
Fomo et al. 1994,<br />
Wilson et al. 1992<br />
Wilson et al. 1992<br />
Kassulke et al.<br />
1990, Wilson &<br />
Flanagan 199 1,<br />
Harley et al. 1985<br />
Julien 1992<br />
Fomo et al. 199 1 b,<br />
Harley et al. 1985,<br />
Kassulke et al. 1990,<br />
Wilson & Flanagan<br />
1991<br />
Fomo et al. 1991b,<br />
Harley et al. 1985<br />
Julien 1992<br />
Julien 1992<br />
Julien 1992<br />
Julien 1992<br />
Australia 1989 Davis et al. 1991,<br />
Wilson & Flanagan<br />
1990<br />
Fomo et al. 1991a,<br />
Julien 1992<br />
Host specificity tests indicate that Phloeospora mimosae-pigrae is specific to M.<br />
pigra and Australian authorities have granted permission to liberate this pathogen.<br />
Testing of Diabole cubensis is still in progress.<br />
Attempts at biological control<br />
AUSTRALIA<br />
M. pigra was probably introduced to Australia at Darwin sometime during the 20 years<br />
before 1891 (Miller and Lonsdale 1987, Lonsdale et al. 1989) and, after a slow start, under-<br />
went a population explosion in the late 1970s and, by 1992, had already covered some 800<br />
km2 of wetlands (Lonsdale 1992). In its century of occupation, at least 114 species of<br />
phytophagous insects have come to attack it (5 Orthoptera, 3 Isoptera, 49 Hemiptera, 21
4.1 6 Mimosa pigra 151<br />
Table 4.16.2 Insects tested against Mimosa pigra, but not released (after Forno 1992).<br />
Species Portion attacked<br />
INSECTS<br />
Coleoptera<br />
CHRYSOMELIDAE<br />
Cryptocephalus (= Diplacaspis) nr<br />
miserabilis leaves<br />
Diplacaspis nr prosternalis stems and leaves<br />
Lexiphanes guerini<br />
young leaves<br />
Syphrea bibiana<br />
seedlings and roots<br />
Lepidoptera<br />
GELECHllDAE<br />
nr Aroga leaves and stems<br />
Table 4.16.3 Natural enemies of Mimosa pigra under investigation (I.W. Forno pers.<br />
comm. 1993).<br />
Species Portion attacked Status<br />
INSECTS<br />
Coleoptera<br />
APlONlDAE<br />
Coelocephalapion pigrae flower buds and leaves<br />
CURCULIONIDAE<br />
Chalcodermus serripes flower buds and immature seeds<br />
FUNGI<br />
Sibinia fastigiata immature seeds<br />
Sibinia ochreosa flower buds<br />
Sibinia peruana flower buds<br />
Sibinia seminicola immature seeds<br />
Diabole cubensis leaves<br />
Phloeospora mimosae-pigra stems, leaves, seed pods<br />
awaiting approval for release<br />
in quarantine in Australia<br />
in quarantine in Australia<br />
under study in Mexico<br />
in quarantine in Australia<br />
in quarantine in Australia<br />
under host testing<br />
approved for release<br />
Coleoptera and 36 Lepidoptera). Of the 11 4,47 species are seldom encountered, 39 are<br />
occasionally found and 28 are common. Thirty of the species are known pests of cultivat-<br />
ed plants and all except two are thought to be polyphagous. These two (a psyllid and a<br />
gelechiid moth) probably feed only on a restricted number of leguminous plants. In this<br />
survey no plant pathogens were recorded (Flanagan et al. 1990, Wilson et al. 1990).<br />
In spite of this diverse insect attack there is still an enormous difference, two orders of<br />
magnitude, between the bank of M. pigra seeds in the soil in Mexico (a mean of 117.5lm2)<br />
and Australia (a mean of 12380lm2). This is believed to reflect the differential occur-<br />
rence of effective natural enemies in each region (Lonsdale and Segura 1987).<br />
The first insects to be liberated-in 1983-for biological control of M. pigra were
152 Biological Control of Weeds: Southeast Asian Prospects<br />
two seed feeding bruchid beetles, Acanthoscelides quadridentatus and A. puniceus, both<br />
of which established readily (Table 4.16.1). They had previously been shown to be ade-<br />
quately host specific (Kassulke et al. 1990). Although these species have become wide-<br />
spread, on average they destroy only 0.8% of mature seed, so are not having much<br />
impact. Of the beetles sampled, 97.8% proved to be A. puniceus (Forno et al. 1991b,<br />
Wilson and Flanagan 1991, Wilson et al. 1992). A parasitoid, Dinarmus sp.<br />
(Pteromalidae), was reared from field-collected bruchids, but did not appear to be having<br />
much influence on beetle populations (C. Wilson pers. comm.).<br />
Next, in 1985, the stem feeding beetle Chlamisus mimosae was released and readily<br />
became established (Forno et al. 1991b) and in 1992193 large populations have been<br />
found at the Finniss R., Northern Territory and are severely damaging M. pigra stems<br />
(I.W. Forno pers. comm. 1 993).<br />
During 1989, two stem boring moths Neurostrota gunniella and Carmenta mimosa<br />
were released. N. gunniella established rapidly and, within a few months, was not only<br />
widespread near the release site but damaging a large number of stems (Forno et al.<br />
1991b, Wilson and Flanagan 1990). By 1993, it had spread to all M. pigra infestations<br />
and is associated with a naturally-occurring, exotic, die-back pathogen (Wilson 1992).<br />
There is a strong negative correlation between seed production and moth populations,<br />
suggesting that N. gunniela can reduce seed numbers by up to 60% (Anon 1992).<br />
Carmenta mimosa is very damaging to young plants and is spreading rapidly in the<br />
Finniss R. region where it is severely damaging stems (I.W. Forno pers. comm. 1993,<br />
Wilson 1992).<br />
The flower bud weevil Coelocephalapion aculeatum was liberated in 1992 and has<br />
become established, but its effects remain to be assessed (Forno et al. 1994, Wilson et al.<br />
1992).<br />
THAILAND<br />
M. pigra was introduced from Indonesia to the Chiang Mai province between 1947 and<br />
the early 1960s as a cover and green manure crop. When found useless for the purpose, it<br />
was employed for the control of irrigation ditchbank erosion, but has now become one of<br />
the worst and most aggressive weeds in the country.<br />
Napompeth (1981) reported 5 insect species attacking M. pigra but, of these, only<br />
the boring beetle Sagra femorata caused much damage. A further study (Napompeth<br />
1983) increased the number to 19 insects, but without adding any promising species.<br />
Both Acanthoscelides puniceus and A. quadridentatus were liberated in 1984 (Table<br />
4.16.1) and are now destroying between 1% and 20% of mature M. pigra seeds, which is<br />
significantly higher than that recorded for Australia (Forno 1992). The reasons for this<br />
different level of effect are not known. Chlamisus mimosae was liberated in 1986 and<br />
became established, but is not causing significant damage. The moth Neurostrota gun-<br />
niella, which is producing such spectacular damage in Australia, has not been liberated in<br />
Thaildnd because it has been shown to be capable of developing in the aquatic Neptunia<br />
oleracea, which is used as a vegetable. Two other species, the weevil Coelocephalapion<br />
aculeatum and the moth Carmenta mimosa were liberated in 1991 (Wilson et al. 1992),<br />
but there is no information on their establishment or impact.
4.1 6 Mimosa pigra 153<br />
MALAYSIA<br />
Acanthoscelides quadridentatus and A puniceus have become established at Kota Bharu<br />
in northern Malaysia adjacent to the region where they are established in Thailand (B.<br />
Napompeth pers. comm. 1993).<br />
MYANMAR<br />
Acanthoscelides quadridentatus and A puniceus have also become established in<br />
Myanmar along the border with Thailand (B. Napompeth pers. comm. 1993).<br />
INDONESIA<br />
M. pigra has been established in Indonesia at least since 1844 but is regarded as a less<br />
serious weed than in Thailand (Napompeth 1982, 1983). At least 10 insects were record-<br />
ed on it and, at times, causing considerable damage in the Bogor area (1 Orthoptera, 3<br />
Hemiptera, 1 Diptera, 2 Coleoptera and 3 Lepidoptera). Only one of these, a cerambycid<br />
borer Milothris irrorata was regarded as having any potential to cause important damage<br />
(Napompeth 1982). As there were doubts about its host specificity (Kalshoven 1981), it<br />
was introduced to Thailand for further study but did not survive in culture (Napompeth<br />
1982, 1992b). Acanthoscelides spp. have been found in M. pigra seed pods collected in<br />
Bogor in 1992, although there are no records of releases having been made (B.<br />
Napompeth pers. comm. 1993).<br />
VIETNAM<br />
Acanthoscelides puniceus and A. quadridentatus from Thailand were liberated in 1987<br />
and Chlamisus mimosae in 1990, but there is no information on their establishment<br />
(Julien 1992).<br />
Important natural enemies<br />
Acanthoscelides quadridentatus Coleoptera: Chrysomelidae<br />
A. puniceus<br />
These two species occur widely in Mexico and A. quadridentatus is also recorded from<br />
Texas, Nicaragua and Honduras. There are also two additional species, A. pigricola and<br />
A. zebratus, that are apparently specific to M. pigra seeds (Habeck and Passoa 1982).<br />
Eggs are laid during the day in or near crevices between pod segments. At 25°C each<br />
larva hatches after 10 days and tunnels through the pod into a single seed in which it<br />
completes its development. Pupation occurs in a cell and adults emerge by chewing a<br />
hole through the seed coat. On average A. quadridentatus females live 93 days and lay 65<br />
eggs, whereas A. puniceus females live 130 days and lay 178 eggs (Kassulke et al. 1990).<br />
Carmenta mimosa Lepidoptera: Sesiidae<br />
This species is native to Mexico and Cuba. In Mexico the larvae tunnel in the stems of<br />
M. pigra, thereby weakening the plant. The upper two-thirds of the stems frequently snap<br />
off, resulting in spectacular damage. Females lay 1 to 5 eggs at a time (up to a total of<br />
about 260) in the axils of the topmost, fully expanded leaves on a stem. At 25°C larvae<br />
hatch after 11 days and tunnel into the stem at a node or the swelling at the base of a leaf<br />
petiole. They feed on the outer layers of the plant, sometimes ringbarking it, or tunnel
154<br />
Biological Control of Weeds: Southeast Asian Prospects<br />
through the pith; they are cannibalistic if they meet another larva. They eject frass onto<br />
the surface of the stem. Occasionally larvae leave the stem and re-enter below the soil<br />
surface, grazing on and in the root, causing damage which sometimes kills the plant.<br />
Depending upon plant quality, there are 8 or 9 larval instars in 40 to 99 days before lar-<br />
vae spin a silken cocoon in which they pupate. The duration of the pupal period is 18 to<br />
21 days, giving an average life cycle of 98 days. Larvae can be reared on an artificial<br />
diet. In host specificity tests, C. mimosa was found to complete its development only on<br />
M. pigra (Forno et al. 1991 a, 1994, Smith and Wilson 1992, Wilson et al. 1992).<br />
Chalcodermus serripes Coleoptera: Curculionidae<br />
This weevil is native to Mexico. Adults feed on young leaves, flower buds and seeds.<br />
Eggs are inserted into the ventral side of pods so that they are at the embryo end of a<br />
developing seed. They hatch in 6 days and larvae feed on the soft developing seed,<br />
destroying the embryo (I.W. Forno, pers. comm.).<br />
Chlamisus mimosae Coleoptera: Chrysomelidae<br />
This species is native to Brazil. Females mate 2 weeks after emergence and then begin to<br />
lay eggs. Each egg is enveloped in faecal material and attached to the underside of a leaf<br />
by a fine stalk. At 25°C larvae hatch after 3 weeks and construct a conical case which is<br />
added to as the larva grows. Larval development time is 83 days and the pupal stage lasts<br />
25 days. Adults live up to 95 days. Adults and larvae graze on the epidermis of the grow-<br />
ing tips, on green stems and on leaves. In Darwin (Northern Territory) cultures of C.<br />
mimosae were attacked by a pupal parasitoid and predatory mites (Wilson et al. 1992).<br />
Coelocephalapion aculeatum Coleoptera: Apionidae<br />
This species is native to Mexico. It lays one egg at a time into a separate flower bud (of<br />
which an inflorescence contains up to 100). Larvae hatch after 2 days and feed on the<br />
developing flower buds, destroying the reproductive parts and sometimes the pedicel.<br />
Larval development takes about 7 days and pupal development 3 days. Preoviposition is<br />
about 7 days, adults live at least 3 months and may lay up to 5 eggs per day. Adults chew<br />
into the unopened flower buds and feed on the anthers and the pistil (Heard 1992, Wilson<br />
et al. 1992). This species could develop satisfactorily only on M, pigra (Forno et al.<br />
1 994).<br />
Coelocephalapion spretissimum and C. pigrae Coleoptera: Apionidae<br />
The life cycle of these species is similar to that of C. aculeatum. Adults feed on young<br />
leaves as well as on flower buds. The host testing of C. pigrae has been completed.<br />
Neurostrota gunnielh Lepidoptera: Gracillariidae<br />
This species is widespread in tropical or subtropical, moderately wet to semi-arid habitats<br />
wherever M. pigra occurs from southern Texas to Costa Rica and Cuba. It has been<br />
established in Australia.<br />
Eggs are laid singly on the ventral side of the first or second leaf from the branch tip<br />
and hatch about 4 days later. The first and second instar larvae are adapted for mining by<br />
being flattened dorso-ventrally, having large blade-like mandibles and no thoracic legs.
4.16 Mimosa pigra 155<br />
Each mines up to 5 leaf pinnules. Third instar larvae are cylindrical, enter the leaf rachis<br />
and tunnel to the stem tip. Later instars usually tunnel down the stem. They sometimes<br />
leave the stem and re-enter it at a node or near a prickle. Frass is usually visible when a<br />
larva is inside a stem. Pupation occurs in a cocoon spun between pinnules or inside the<br />
stem. The outside of the cocoon is ornamented with small, pearly-white, frothy balls dis-<br />
charged from the anus.<br />
At 25°C the time from egg to adult is about 30 days and equal numbers of males<br />
and females are produced. Females mate on the night of emergence and lay an average of<br />
86 eggs, most on the second night. N. gunniella caused very severe damage to M. pigra<br />
in quarantine trials which demonstrated that, except for attack on Neptunia species, it<br />
was specific to M. pigra. The aquatic Neptunia oleracea, which is an important vegetable<br />
in Thailand was attacked, so it has not been released there. In Mexico N. gunniella larvae<br />
are heavily attacked by parasitoids (Davis et al. 1991, Forno et al. 1989a, 199 1 b).<br />
Sibinia spp. Coleoptera: Curculionidae<br />
S. fastigiata occurs from Mexico to Brazil and Peru, whereas S. seminicola occurs from<br />
Texas and Mexico to southeastern Brazil and northeastern Argentina. These two Sibinia<br />
species are larger than the two that follow. Larvae of both species develop in the pods of<br />
M. pigra and feed on the seeds. Larvae of S. seminicola feed on green, immature seeds<br />
and pupation occurs within the pods while they are still attached to the plants (Clark<br />
1984).<br />
Sibinia ochreosa occurs in Texas, Mexico, Honduras, Nicaragua, Brazil, Argentina<br />
and S. peruana occurs in Mexico, Guatemala, Honduras, Costa Rica, Panama, Brazil,<br />
Peru, Bolivia and Argentina. Larvae of these two species develop in the flower buds of<br />
M. pigra and, at least the former species, pupates in the flower head (Clark 1984).<br />
FUNGI<br />
Diabole cubensis Fungi: Uredinales<br />
This rust attacks the leaves of M. pigra in Mexico and Cuba. It is particularly common<br />
and damaging during the dry season when there are high day temperatures and a signifi-<br />
cant drop at night leading to dew formation. Five hyperparasitic fungi are consistently<br />
encountered, often completely overgrowing the rust (Evans 1988, 1990).<br />
Phloeospora mimosae-pigrae Fungi: Coelomycetes<br />
This fungus causes extensive defoliation during the wet season, particularly in the Gulf<br />
coast of Mexico, but also occurs in Trinidad, Venezuela, Colombia and Brazil. It attacks<br />
stems, leaves and seed pods. It is host specific to M. pigra (Evans 1988, 1990, Seier and<br />
Evans 1993).<br />
Comment<br />
The majority of natural enemies of M. pigra so far studied are flower or seed attacking<br />
insects and there are indications already that cbnsiderable amounts of seed are being<br />
destroyed-up to 60% from Neurostrota alone in Australia and up to 20% by bruchids in
156<br />
Biological Control of Weeds: Southeast Asian Prospects<br />
Thailand. However, starting from a seed bank of 9000 per m2, this degree of reduction is<br />
nowhere like limiting.<br />
The inhibition to tip growth produced in Australia by Neurostrota gunniella is likely<br />
to be far more significant and will become even more so if the borer, Carmenta mimosa,<br />
becomes abundant enough to weaken a considerable proportion of larger stems.<br />
If (when) the two apparently specific fungi are approved for release, it is confidently<br />
expected that they will make a major contribution, Phloeospora mimosa-pigrae in the<br />
wet season and the rust Diabole cubensis in the dry.<br />
It is highly probable that a complex of natural enemies will be required to bring<br />
about an adequate reduction in competitiveness of M. pigra. It is still unclear whether<br />
those available or under investigation will be adequate but, if required, there are addition-<br />
al species that could be examined and future priority might well be given to leaf, stem or<br />
root feeding species.
158<br />
Biological Control of Weeds: Southeast Asian Prospects<br />
Mimosa pudica<br />
(after Holm et a/. 1977)
Map 4.17 Mimosa pudica<br />
4.17 Mimosa pudica<br />
Mimosa pudica<br />
Very little is known about the natural enemies of Mimosa pudica in its centre of origin in<br />
Central America. Most of the species recorded from there or elsewhere are widely<br />
polyphagous and few show promise as biological control agents. Since at least three<br />
forms of the weed are known it would be desirable to establish which forms are weedy in<br />
Southeast Asia so as to enable any searches in Central America to concentrate on that<br />
form. On general grounds, it is probable that useful species do exist.
160 Biological Control of Weeds: Southeast Asian Prospects<br />
4.17 Mimosa pudica L.<br />
Mimosaceae<br />
common sensitive plant; paklab, sampeas (Cambodia), daoen kaget kaget<br />
(Indonesia) mala malu (Malaysia), makahiya (Philippines), mai yarap (Thailand)<br />
mac co (Vietnam)<br />
Rating<br />
+++ Myan, Sing, Indo<br />
17 ++ Msia, Brun, Phil<br />
+ Thai, Viet<br />
. Laos, Camb<br />
Origin<br />
Tropical America.<br />
Distribution<br />
M. pudica is widespread in tropical, subtropical and temperate areas of the world. Its dis-<br />
tribution and other relevant aspects were summarised by Waterhouse and Norris (1987).<br />
There are at least three distinct varieties (Brenan 1959). M. pudica hispida is uncommon<br />
in the Americas, but is established in the Philippines (Bameby 1989), the Caroline and<br />
Mariana Is, Queensland, India and in African savanna country. M. pudica unijuga occurs<br />
in Hawaii and probably in other Pacific countries where it is a major weed.<br />
Characteristics<br />
M. pudica is low, much branched, generally perennial, slightly woody at the base, from<br />
15 to 100 cm high and has either an upright or a low trailing habit. Its stiff reddish-brown<br />
or purple stems bear scattered thorns. The hairy leaves are alternate, bipinnate and com-<br />
pound. They are sensitive to the touch, the petiole dropping and the leaflets being rapidly<br />
drawn back and folded. The pink flowers form small globular heads, each borne on a<br />
short hairy stalk arising from a leaf axil. Seeds are produced in pods which split into sin-<br />
gle-seeded segments bearing bristles, which aid dispersal by animals. In tropical coun-<br />
tries the weed flowers all year and each plant may produce up to 700 seeds.<br />
Importance<br />
M. pudica is a weed in 22 crops in 38 countries (Holm et al. 1977). It is common in waste<br />
land and is also a weed of lawns, crops, pastures and roadsides. In Southeast Asia and the<br />
Pacific it is a serious weed in maize, sorghum, sugarcane, tea, soybeans, upland rice,<br />
pinea~jples and cotton. Because of its tolerance to shading it is an important weed in plantation<br />
crops, such as rubber, coconuts, bananas, papaya, coffee, oil palm and citrus. In<br />
tropical pastures its dense growth and thorns often deter animals from feeding on suitable<br />
forage mingled with it (Holm et al. 1977). The thorns deter hand weeding and, as it sur-
4.1 7 Mimosa pudica 161<br />
vives mowing, it is a very unwelcome component of lawns. Attempts to select thornless<br />
types as pasture plants have not been successful.<br />
Natural enemies<br />
Some information is summarised by Waterhouse and Norris (1987). It is interesting that<br />
Holm et al. (1977) report M. pudica to be a widespread weed in the Caribbean, but far<br />
less important to the north and south of this region. This suggests that it evolved else-<br />
where in the Americas and has not been accompanied into the Caribbean by its full suite<br />
of natural enemies. Nevertheless a preliminary survey in Trinidad (Table 4.17.1) revealed<br />
14 insects attacking it, but they are probably polyphagous, with the possible exception of<br />
the arctiid caterpillar Lophocampa catenulata and the beetle, Chlarnisus sp. (Yaseen<br />
197 1, 1972). Perez et al. (1 988) found that the race Jilenus of Hemiargus hanno in Cuba<br />
appears not to attack plants other than M. pudica, although partially grown larvae of the<br />
Trinidad race were able to complete their development on Aeschynomene sensitiva and<br />
Cajanus cajan (Yaseen 1972). H. hanno Jilenus feeds readily on M. pudica seeds and is<br />
particularly active in spring when the weed is producing most seed (Perez et al. 1988).<br />
Four additional insects are known from Brazil (Garcia 1982a,b, 1983) but, so far, no<br />
species of Heteropsylla, although a special search for one might be rewarding. Although<br />
M. pudica was often encountered in surveys carried out in Mexico and Venezuela for nat-<br />
ural enemies of M. pigra, casual observation did not suggest that it was heavily attacked,<br />
less so indeed than M. invisa (I.W. Forno pers. comm. 1993).<br />
It is not known whether there is any differential attack by natural enemies on any of<br />
the three or more varieties of M. pudica, which have been established on morphological<br />
differences alone.<br />
Table 4.17.1 Natural enemies of Mimosa pudica.<br />
Species Country Portion of plant attacked References<br />
INSECTS<br />
Hemiptera<br />
COCCIDAE<br />
Coccus longulus Fiji<br />
CYDNIDAE<br />
Microporus<br />
(= Microcompsus) sp. Trinidad<br />
DlASPlDlDAE<br />
Hemiberlesia lataniae Fiji<br />
Pinnaspis strachani Fiji<br />
MARGARODIDAE<br />
Icerya seychellarurn Fiji<br />
SCUTELLERIDAE<br />
2 species Trinidad<br />
Coleoptera<br />
CHRYSOMELIDAE<br />
Chlarnisus sp. Trinidad<br />
stems Hinckley 1963<br />
stems<br />
stems<br />
stems<br />
flowers<br />
Yaseen 1972<br />
Hinckley 1963<br />
Hinckley 1963<br />
Hinckley 1963<br />
Yaseen 197 1<br />
flower buds Yaseen 197 1, 1972<br />
(continued on next page)
162 Biological Control of Weeds: Southeast Asian Prospects<br />
Table 4.17.1 (continued)<br />
Species<br />
CURCULIONIDAE<br />
Chalcodermus sp.<br />
Promecops<br />
?campanulicollis<br />
Country Portion of plant attacked References<br />
Brazil<br />
Brazil<br />
seed pods<br />
leaves; mainly on<br />
M. invisa, see table<br />
4.15.1<br />
C.A. Garcia pers. comm.<br />
Garcia 1982a,b, 1983<br />
An unidentified sp. Trinidad Yaseen 197 1<br />
Lepidoptera<br />
ARCTllDAE<br />
Lophocampa catenulala<br />
GEOMETRIDAE<br />
Ptychamalia perlata<br />
GRACllARllDAE<br />
Neurostrota gunniella<br />
HESPERIIDAE<br />
Cogia (= Caicella)<br />
calchas (= ?<br />
Nisoniades bessus)<br />
?Slaphylus mazans<br />
LYCAENIDAE<br />
Calephelis sp.<br />
Hemiargus hanno<br />
Trinidad leaves Yaseen 1971<br />
Trinidad leaves<br />
Mexico leaves, stems<br />
Trinidad leaves, flowers<br />
Yaseen 1972<br />
Davis et al. 1991<br />
Cock 1985, Yaseen 1972<br />
Trinidad leaves, flowers Yaseen 1972<br />
Trinidad<br />
Cuba,<br />
Trinidad<br />
leaves<br />
leaves, flowers, pods<br />
also on flowers of<br />
Aeschynomene<br />
semitiva and Cajanus<br />
cajan (= C. indicus)<br />
Tmolus azia Trinidad leaves, flowers Yaseen 1972<br />
NOCTUI DAE<br />
Spodoptera litura Fiji<br />
Sp. 1 (velvet black larva) Brazil<br />
PlERlDAE<br />
Eurema lisa Cuba<br />
Eurema tenella Brazil<br />
also on Mimosa pudica<br />
and groundnut flowers<br />
leaves<br />
leaves; also on<br />
M. invisa<br />
leaves: mainly on<br />
M. invisa, see table<br />
4.15.1<br />
Yaseen 1971<br />
Dethier 1940, Perez et al.<br />
1988, Yaseen 1972<br />
Hinckley 1963<br />
Garcia 1982a,b, 1983<br />
Dethier 1940<br />
Garcia 1982a,b, 1983<br />
TORTRICIDAE<br />
Platynota rostrana Trinidad leaves Yaseen 1972<br />
NEMATODE<br />
Meloidogyne sp. Cuba Holm et al. 1977,<br />
Izquierdo et al. 1987<br />
FUNGI<br />
VIRUS<br />
Oidium sp. Mexico<br />
?Puccinia sp. Mexico<br />
mildew on leaves<br />
rust on leaves<br />
Evans 1987<br />
Evans 1987<br />
unspecified Germany Umrath et al. 1979
164 Biological Control of Weeds: Southeast Asian Prospects<br />
Monochoria vaginalis<br />
(after Holm et a/. 1977)
Map 4.18 Monochoria vaginalis<br />
4.1 8 Monochoria vaginalis 165<br />
Monochoria vaginalis<br />
Monochoria vaginalis appears to be a major weed only in Southeast Asia and then only<br />
in rice. Almost nothing is known of its natural enemies in India and Africa where it<br />
occurs, but is not regarded as important. This suggests that a survey in these regions<br />
might reveal promising biological control agents.
166 Biological Control of Weeds: Southeast Asian Prospects<br />
4.18 Monochoria vaginalis (Burm. f.) Presl<br />
Pontederiaceae<br />
monochoria; ka kiad chrach (Cambodia), phak kbiat (Thailand), rau mac la thon<br />
(Vietnam), etjeng padi (Indonesia), biga bigaan (Philippines), kelayar, echeng<br />
padi (Malaysia)<br />
Rating<br />
+++ Laos, Camb, Msia, Brun, Indo, Phil<br />
26 ++ Myan, Thai, Viet, Sing<br />
Origin<br />
Tropical Asia and Africa (Holm et al. 1977), but not a pest in Africa or Asia (Soerjani et<br />
al. 1987), although it is clearly very important in Southeast Asia.<br />
Distribution<br />
Africa, India, China, Korea, Japan, Southeast Asia to northern Australia, Fiji and Hawaii.<br />
Characteristics<br />
A smooth, fleshy, semi-aquatic annual or perennial, 0.1 to 0.5 m tall; the plant roots in<br />
mud and its upper portions grow above water; stemless, base of leaves heartshaped or<br />
rounded, shiny deep green; petioles soft, hollow; inflorescence with a large bract and<br />
arising about two thirds of the way up the petiole from the base and opposite the leaf; 3<br />
to 25 violet or lilac flowers producing numerous, small seeds throughout most of the<br />
year. Seed germination and seedling growth not reduced by submergence. Old plants<br />
often form large clumps.<br />
Importance<br />
M. vaginalis occurs in marshy places, freshwater pools, mudflats, ditches, along canal<br />
banks and in rice fields. It is a very serious weed of rice in eastern and southern Asia. It<br />
is predominantly an annual in flooded ricefields, dying when the fields dry out, but<br />
developing again later from seed.<br />
In Taiwan, M. vaginalis produced more fresh tissue weight than any other weed in<br />
rice, twice that of second ranking Echinochloa crus-galli (Lin 1968). However in the<br />
Philippines it was outcompeted in rice by E. crus-galli (Lubigan and Vega 1971). Only in<br />
rice is it reported as a very widespread and important weed, except for its occurrence in<br />
taro in Hawaii.<br />
Its leaves are eaten as a pot herb in India and several parts of it are used as herbal<br />
medicine, the juice being prescribed for various conditions and the roots for stomach and<br />
liver ailments and toothache (Burkill 1935, Soerjani et al. 1987).
Natural enemies<br />
4.1 8 Monochoria vaginalis 167<br />
So little is recorded about the natural enemies of M. vaginalis (Table 4.18.1) that it is not<br />
possible to assess the prospects for biological control. However, the fact that it is appar-<br />
ently not a weed in Africa or western Asia suggests that it would be well worthwhile<br />
investigating these regions for suitable agents.<br />
Table 4.18.1 Natural enemies of Monochoria vaginalis.<br />
Species Location Other hosts References<br />
INSECTS<br />
Orthoptera<br />
ACRlDlDAE<br />
Gesonula punctifrons India<br />
Hemiptera<br />
CICADELLIDAE<br />
Macrosteles fasciffons USA rice<br />
DELPHACIDAE<br />
Tarophagus proserpina Philippines taro, cassava, sweet potato<br />
Lepidoptera<br />
NOCTUIDAE<br />
Spodoptera litura India widely polyphagous<br />
PYRALIDAE<br />
Elophila responsalis Indonesia Marsilea minuta, Pistia<br />
stratiotes, Salvinia molesta,<br />
S. cucullata<br />
Nymphulafregonalis India probably polyphagous<br />
SPHlNGlDAE<br />
Hippotion echeclus India polyphagous<br />
NEMATODES<br />
Hirschmaniella spp. rice, sugarcane,<br />
many weeds<br />
Meloidogyne graminicola rice, many weeds<br />
VIRUSES<br />
rice ragged stunt Thailand, rice<br />
Philippines<br />
Pistia virus India<br />
FUNGI<br />
Cercospora sp. India<br />
Doassansia sp. India<br />
Rhizoctonia solani India rice, potato<br />
Thanatephorus cucumeris Philippines rice<br />
a leaf blight Philippines Sphenoclea zeylanica<br />
CRUSTACEA<br />
Triops cancriformis Japan Veronica peregrina and<br />
some other weeds<br />
Sankaran & Rao 1972<br />
Way et al. 1983<br />
Duatin & Pedro 1986<br />
Sankaran & Rao 1972<br />
Handayani & Syed 1976,<br />
Mangodihardjo 1975<br />
Sankaran & Rao 1972<br />
Sankaran & Rao 1972<br />
Luc et al. 1990<br />
Luc et al. 1990<br />
Parejarearn et al. 1988,<br />
Salamat et al. 1987<br />
Menon & Ponnappa 1964<br />
Menon & Ponnappa 1964<br />
Menon & Ponnappa 1964<br />
Gokulapalan & Nair 1983<br />
Moody et al. 1987,<br />
Mew et al. 1980<br />
Bayot et a]. 1992<br />
Igarashi 1985
168 Biological Control of Weeds: Southeast Asian Prospects<br />
Nep hrolepis biserra ta<br />
(after Barnes and Chan, 1990)
Map 4.19 Nephrolepis biserrata<br />
4.19 Nephrolepis biserrata 169<br />
Nephrolepis biserrata<br />
N. biserrata is a widespread fern that probably originated in Tropical Africa. Almost<br />
nothing was learnt concerning natural enemies from a literature search. A survey in its<br />
area of origin would be required to evaluate whether there were any natural enemies that<br />
might be of value for biological control.
170 Biological Control of Weeds: Southeast Asian Prospects<br />
4.19 Nephrolepis biserrata (Sw.) Schott<br />
Nephrolepidaceae (formerly in the Davalliaceae)<br />
broad sword fern; paku larat (Malaysia)<br />
Rating<br />
+++ Sing, Brun<br />
10 ++ Msia, Phil<br />
Thailand<br />
Origin<br />
Probably Tropical Africa.<br />
Distribution<br />
Pantropical. N. biserrata occurs throughout tropical Africa, ranging in the west from<br />
Guinea to Angola and in the east from Sudan to Durban (South Africa). It is mainly<br />
coastal, but infrequent in the interior. It is most abundant up to 350 m (Jacobsen 1983). In<br />
addition to Southeast Asia, it is also known from India, Australia, Japan, the Pacific,<br />
USA and Mexico.<br />
Characteristics<br />
N. biserrata is a perennial, terrestrial or epiphytic fern. The rhizome bears abundant<br />
scales and produces many long stolons. The fronds are tufted, suberect to arching and<br />
green when young, turning brown when old. The pinnae are commonly 15 to 20 cm<br />
wide, exceptionally more than 30 cm. Fertile pinnae are narrower than the sterile pinnae<br />
and bear sori well clear of the edge. The veins are indistinct and fork once or twice.<br />
When rooted in the soil, the fern commonly reaches a height of 2m and, exceptionally in<br />
South Africa, up to 4 m.<br />
Importance<br />
N. biserrata is very common in shaded places in the lowlands wherever the conditions<br />
are not too dry. It can form dense masses in rubber and oil palm plantations and in<br />
orchards and often occurs as an epiphyte on palms. Where pineapples had been grown<br />
for up to 10 years in Malaysia, 90% of the viable seeds and spores in the top 15 cm of<br />
soil were spores of N. biserrata (in particular) and 8 other ferns (Wee 1974), enabling its<br />
rapid reappearance after cultivation.<br />
Natural enemies<br />
The only records of natural enemies encountered were those of an eriophyid mite on<br />
N. biserrata in Fiji (Mani and Jayaraman (1987) and of the nematode Aphelenchoides<br />
fragariae in Hawaii, but there is no evidence that a careful search has ever been made.
Attempts at biological control<br />
There have been none.<br />
Comment<br />
4.19 Nephrolepis biserrata 171<br />
The genus Nephrolepis contains about 20 species (Tryon and Tryon 1982), or 35 species<br />
(Jacobsen 1983). It is primitive among the group of oleandroid, davallioid and nephrole-<br />
poid ferns to which it belongs and N. biserrata and its close allies represents the more<br />
primitive element in the genus Nephrolepis (Nayar and Bajpai 1976). A survey for natur-<br />
al enemies, particularly in the areas in Africa where it occurs, would reveal whether there<br />
are any that might be of value in biological control.
172 Biological Control of Weeds: Southeast Asian Prospects<br />
Panicum repens<br />
(after Holm et a/. 1977)
Map 4.20 Panicum repens<br />
4.20 Panicum repens<br />
Panicum repens<br />
P. repens is a major grassy weed in Southeast Asia. It is probably of tropical<br />
AfricanJMediterranean origin, a region where it is not reported to be a problem. Very few<br />
natural enemies have been recorded and, without a preliminary survey in its area of ori-<br />
gin, it would not be possible to evaluate the prospects for its biological control.
174 Biological Control of Weeds: Southeast Asian Prospects<br />
4.20 Panicum repens L.<br />
Poaceae<br />
torpedo grass, creeping panic grass; myet kha (Myanmar) yah chan ah kat, yah<br />
chanagard (Thailand), chhlong (Cambodia), co 6ng (Vietnam), keruong padi,<br />
rumput jae jae (Malaysia and Indonesia), luya luyahan (Philippines)<br />
Rating<br />
+++ Viet, Indo<br />
16 ++ Thai, Msia, Sing, Phil<br />
+ Myan, Brun<br />
Laos. Camb<br />
Origin<br />
Tropical and North Africa, Mediterranean (sometimes said to be native to Asia).<br />
Distribution<br />
Panicum repens is widely distributed in the tropics and subtropics. It was introduced to<br />
Java about 1850, but is said not to occur in the Moluccas. No seeds are produced in<br />
Indonesia (Soerjani et al. 1987).<br />
Description<br />
Panicum repens is an erect, wiry, creeping, perennial grass, rooting at hairless nodes and<br />
bearing flowering stalks 30 to 90 cm tall. It spreads widely (up to 7 m), but does not form<br />
dense clumps. Its smooth, sharp-pointed, branched rhizomes are often swollen or knotty<br />
and have brownish or whitish scales. Its leaves are alternate. The inflorescence is an open<br />
terminal panicle, 6 to 20 cm long, with many tender branches pointing obliquely<br />
upwards. The spikelets are pale green or pale yellow and often tinged with purple.<br />
Importance<br />
Panicum repens is one of the most serious grass weeds because of its rapid rate of spread<br />
and the persistence and hardiness of its coarse, enlarged rhizomes. It suppresses other<br />
plants by its allelopathic (inhibiting) products (Perera et al. 1989). It is primarily a weed<br />
of moist, coastal, sandy soils, although it also grows in heavy upland soils (to 2000 m in<br />
Indonesia). It thrives in open sunny situations, but can stand partial shade and its rhi-<br />
zomes survive even prolonged dry periods. It tolerates temporary flooding, sometimes<br />
forms floating mats, and encroaches upon ditches, drains and watercourses. It is common<br />
in cultivated lands, grasslands, roadsides and gardens and is frequently reported as a weed<br />
in lawns. In Malaysia it is a serious weed of cocoa, coconuts and rubber, in Indonesia of<br />
rainfed and upland rice, cocoa, coconut, maize, rubber, sugarcane and tea and in Thailand<br />
of rice and orchard crops. In improved pastures it chokes out more nutritionally valuable
4.20 Panicum repens 175<br />
grasses. Deep ploughing favours its spread by breaking up and dispersing its rhizomes.<br />
P. repens is quite palatable to stock when young and has the advantage of being able<br />
to stand heavy grazing and trampling. However, it contains only 3.3% crude protein and<br />
up to 39% crude fibre, so there are other far more nutritious grasses suitable for the same<br />
environment (Holm et al. 1977).<br />
Natural enemies<br />
As can be seen in table 4.20.1, very few natural enemies were revealed by an extensive<br />
literature search. The only species not known to be both polyphagous and a pest is the<br />
mite Parasteneotarsonemus panici, recorded so far only from Tamil Nadu, India,<br />
where it was found causing rusting symptoms beneath the leaf sheath of P. repens<br />
(Mohanasundaram 1984).<br />
Absence of attack almost certainly does not represent the true situation, but rather<br />
that no relevant surveys have been carried out. For example, there has been little advance<br />
in knowledge of cecidomyiid gall flies attacking Panicum spp. since the summary by<br />
Barnes (1954b) in which were listed at least 13 species (Table 4.20.2). Two of these (the<br />
widespread rice stem gall midge Orseolia (= Pachydiplosis) oryzae and Contarinia<br />
(= Stenodiplosis) panici are known to be pests (Hegdekatti 1927). The scarcity of infor-<br />
mation from Africa points to an important gap in our knowledge and the absence of<br />
records from the Mediterranean, where most plants have been studied in some detail for<br />
native-insects, suggests that P. repens is not native to that region.<br />
Comment<br />
Clearly insufficient information is known about the natural enemies of P. repens<br />
(although it doubtless must have many) to provide any meaningful assessment of the<br />
prospects for its biological control. However, the genus Panicum contains a number of<br />
good to very good fodder species (e.g. P. antidotale (blue panic), P. bulbosum (bulbous<br />
panic), P. capillare (witchgrass), P. maximum (guinea grass), P. paludosum (swamp<br />
panic)) as well as several that are weedy and may cause photosensitivity or poisoning<br />
(e.g. P. coloratum (coolah grass), P. luzonense, P. miliaceum (millet panic, or proso), P.<br />
novemnerve). There are thus likely to be considerable problems in discovering organisms<br />
of adequate specificity. Nevertheless, P. repens is not reported as a weed of crops in trop-<br />
ical Africa or the Mediterranean (Holm et al. 1977) and a preliminary survey there might<br />
well reveal promising natural enemies.
176 Biological Control of Weeds: Southeast Asian Prospects<br />
Table 4.20.1 Natural enemies of Panicum repens.<br />
Natural enemies Recorded from References<br />
INSECTS<br />
Hemiptera<br />
CICADELLIDAE<br />
Thaia oryzivora Thailand<br />
DELPHACIDAE<br />
Delphacodes idonea USA<br />
Sogatella kolophon Australia, Pacific Is, SE Asia,<br />
USA, Central & S. America,<br />
W. Africa<br />
Lepidoptera<br />
EUPTEROTIDAE<br />
Nisaga simplex India<br />
NOCTUl DAE<br />
Sesamia cretica Egypt<br />
PYRALI DAE<br />
Cnaphalocrocis medinalis Philippines<br />
Cnaphalocrocis (= Marasmia) Philippines<br />
patnalis<br />
Parapoynx stagnalis<br />
(= Nymphula depunctalis) India<br />
MITES<br />
TARSONEMIDAE<br />
Parasteneotarsonemus panici India<br />
FUNGI<br />
Claviceps sp.<br />
Pyricularia sp.<br />
Pyricularia oryzae<br />
Sporisorium overeemi<br />
NEMATODE<br />
Meloidogyne graminicola<br />
India<br />
India<br />
Leeuwangh & Leuamsang 1967<br />
Ballou et al. 1987<br />
Ballou et al. 1987<br />
Patnaik et al. 1987<br />
Ahmed 1980<br />
Abenes & Khan 1990<br />
Abenes & Khan 1990<br />
Pillai & Nair 1979<br />
Mohanasundaram 1984<br />
Janardhanan et al. 1982<br />
Hilda & Suranarayanan 1976,<br />
Holm et al. 1977<br />
Paje et al. 1964<br />
Rifai 1980<br />
Luc et al. 1990
4.20 Panicum repens 177<br />
Table 4.20.2 Gall flies (Cecidomyiidae) reported attacking Panicum spp. (after<br />
Barnes 1954b).<br />
Species Recorded from Location<br />
Contarinia (= Srenodiplosis) panici<br />
Lasioptera (= Dyodiplosis)fluitans<br />
Lasioptera inustorum<br />
Lasioptera kanni<br />
Lasioptera panici<br />
Lasioptera paniculi<br />
Orseolia cynodontis<br />
Orseolia (= Courteia) graminis<br />
Orseolia (= Dyodiplosis) andropoginis<br />
Orseolia (= Dyodiplosis) fluvialis<br />
Orseolia (= Pachydiplosis) oryzae<br />
Parallelodiplosis javanica<br />
Parallelodiplosis spp.<br />
Yugoslavia, USSR<br />
Panicum fluitans S. India<br />
USA<br />
S. India<br />
USA<br />
Philippines<br />
France, Italy, Algeria, Eritrea,<br />
Senegal<br />
Java, Sri Lanka, S. India<br />
Panicum jluitans S. India<br />
S and SE Asia, W. Africa<br />
Panicum indicum Sri Lanka<br />
Middle East, Java, Peru
178 Biological Control of Weeds: Southeast Asian Prospects<br />
Paspalurn conjuga turn<br />
(after Holm et a/. 1977)
Map 4.21 Paspalurn conjugaturn<br />
4.2 1 Paspalurn conjugaturn 179<br />
Paspalurn conjugaturn<br />
P. conjugaturn is of Tropical American origin, but it is recorded as an important weed in a<br />
number of situations in the Caribbean area. Very few natural enemies have been reported.<br />
A preliminary survey would be required in its centre of origin before the prospects for its<br />
biological control could be evaluated.
180 Biological Control of Weeds: Southeast Asian Prospects<br />
4.21 Paspalurn conjugaturn Bergius<br />
Poaceae<br />
sourgrass; paitan, rumput canggah, rumput pait (Indonesian), rumput kerbau,<br />
jampang canggah, buffalo grass (Malaysia), hulape (Philippines), ya hep<br />
(Thailand)<br />
Rating<br />
+++ Viet, Msia, Indo<br />
15 ++ Sing, Brunei, Phil<br />
Myan, Thai, Laos, Camb<br />
Origin<br />
Tropical America.<br />
Distribution<br />
Paspalurn conjugaturn occurs as a troublesome weed in Central America, West Africa<br />
and the islands and peninsulas of Southeast Asia and the Pacific. These are, for the most<br />
part, the humid tropics (Holm et al. 1977).<br />
Characteristics<br />
P. conjugaturn is a creeping, stoloniferous, perennial grass. The stolons are up to 2 m in<br />
length, often reddish purple in colour and bear roots and a tuft of green to purple leaves<br />
at each node.<br />
The flower stalks are erect, range up to 60 cm, and have smooth nodes. At the apex<br />
of each stalk there are two racemes (flower spikes) 4 to 15 cm long. The stigmas are<br />
white and the anthers bright yellow. The weed can be recognised when in flower by the<br />
typical T-shaped inflorescence.<br />
Importance<br />
P. conjugaturn is mainly a weed of the warm, wet lowlands, although in Hawaii and Sri<br />
Lanka it grows up to 1875 m. It is found in waste areas and along paths and streams, its<br />
inflorescences trailing in the water. It is common in cultivated fields and in natural and<br />
poorly managed pastures and particularly in perennial or plantation crops where the soil<br />
is not ploughed frequently. It spreads rapidly by its stolons, forming dense masses which<br />
can suppress or eliminate tree seedlings and other small plants. It tolerates some shade<br />
and can grow on poor and acid soils.<br />
In the Philippines it flowers all year round and one plant can produce 1500 seeds. It<br />
is also dispersed by broken pieces of stolons rooting after being spread by machines used<br />
for cultivation. In the Philippines it is particularly important in bananas, coffee, papaya,<br />
rice and pineapple; in Cambodia in rice; in Malaysia in citrus, coconuts, oil palm, rice
4.21 Paspalurn conjugaturn 181<br />
and rubber; in Indonesia in tea, oil palm and rubber; and elsewhere in cassava, cocoa,<br />
lawns, maize, pastures, sugarcane and vegetables.<br />
P. conjugaturn is suitable for grazing only when young and the seeds of older plants<br />
have been reported to choke animals by sticking in their throats (Holm et al. 1977).<br />
Natural enemies<br />
The natural enemies of P. conjugatum that have been recorded in the literature (Table<br />
4.21.1) are almost all polyphagous and many of them are of economic importance. One<br />
possible exception is the bagworm moth Brachycyttarus griseus, which was originally<br />
described from Vietnam and is also recorded from Malaysia and the Philippines, as well<br />
as from Guam and Hawaii to which it has spread. It feeds on P. conjugaturn in Hawaii<br />
and on the grass Zoysia pungens in Guam: it probably also feeds on other grasses. In<br />
Guam it is parasitised by the tachinid fly, Stornatomyia sp. (Davis 1990). However, it<br />
does little to control P. conjugaturn in the countries where it already occurs, so it does not<br />
appear to be a promising species to introduce elsewhere.<br />
Perhaps more valuable is the cecidomyiid fly Cleitodiplosis grarninis, a gall forming<br />
fly described from Brazil. The gall consists of the terminal leaves becoming clustered<br />
into an ovoid 30 x 20mm mass as a result of the upper internodes being greatly reduced.<br />
Thirty to forty sulphur-coloured larvae may be found in a single gall, in which they<br />
pupate. They appear in August and September (Barnes 1956). Barnes (1954b) comment-<br />
ed that no gall midge had, at that time, been recorded from the inflorescence of<br />
Paspalurn, although Parallelodiplosis paspali (from Java and India) and Lasioptera sp.<br />
had been recorded from the stems.<br />
The chrysomelid beetle Colaspis (= Maecolaspis) aerea also damages cocoa in<br />
Brazil (Fenonatto 1986) and hence would not be acceptable for introduction elsewhere.<br />
The seeds of P. conjugaturn are harvested by ants in some areas, but the impact of<br />
this on weed density in the field has not been established. Under experimental field con-<br />
ditions in Mexico, the pestiferous fire ant, Solenopsis gerninata, reduced P. conjugaturn<br />
seed densities by 97% or more but had no effect on the seed densities of Bidens pilosa<br />
(Carroll and Risch 1984).<br />
Comment<br />
As with Panicurn repens, so little is known about the natural enemies of Paspalurn conju-<br />
gatum that it is not possible to evaluate the prospects for its biological control. Again,<br />
there is the limitation that some closely related species in the genus Paspalum are of eco-<br />
nomic value for fodder (e.g. P. dilataturn (paspalum), P. distichurn (saltwater couch), P.<br />
plicatulum, P. scrobiculaturn (scrobic), P. vaginaturn (saltwater couch)). However, a fun-<br />
gus attacks the seeds of the first two species, producing the toxin, ergot, and most of the<br />
species are considered weedy in at least some situations, so there may be occasions<br />
where a conflict of interests would have to be resolved, should effective agents be dis-<br />
covered. The prospects for finding these are not perhaps very promising, since Holm et<br />
al. (1977) record P. conjugaturn as a weed in a number of countries in and around the<br />
Caribbean.
182 Biological Control of Weeds: Southeast Asian Prospects<br />
Table 4.21.1 Natural enemies of Paspalum conjugatum.<br />
Species Country References<br />
INSECTS<br />
Thysanoptera<br />
Haplothrips gowdeyi Hawaii Sakimura 1937<br />
Haplothrips paumalui Hawaii Sakimura 1937<br />
Coleoptera<br />
CHRYSOMELIDAE<br />
Colaspis (= Maecolaspis) aerea Brazil<br />
Diptera<br />
CEClDOMYllDAE<br />
Cleitodiplosis graminis Brazil<br />
Lepidoptera<br />
ARCTllDAE<br />
Creatonotos (= Amsacta) gangis<br />
HESPERllDAE<br />
Taractrocera ina<br />
PSYCHIDAE<br />
Brachycyttarus griseus<br />
FUNGI<br />
PYRALIDAE<br />
Cnaphalocrocis medinalis<br />
Cnaphalocrocis (= Marasmia)<br />
patnalis<br />
Parapoynx stagnalis<br />
(= Nymphula depunctalis)<br />
Exserohilum paspali<br />
Leptosphaeria proteispora<br />
Myriogenospora atramentosa<br />
Physarum cinereum<br />
Sorosporium paspali<br />
BACTERIA<br />
Xanthomonas albilineans<br />
Ferronatto 1986<br />
Barnes 1956<br />
Philippines Catindig et al. 1993<br />
Australia Common & Waterhouse 198 1<br />
Guam, Hawaii, Malaysia, Davis 1990<br />
Vietnam, Philippines<br />
Philippines Abenes & Khan 1990<br />
Philippines Abenes & Khan 1990<br />
Philippines Bandong & Litsinger 1984<br />
Brazil Muchovej & Nesio 1987<br />
Hawaii Stevens 1925<br />
Brazil, USA, Venezuela Hanlin & Tortolero 1990<br />
Brazil Muchovej & Muchovej 1987<br />
Hawaii Stevens 1925<br />
Australia Persley 1973<br />
NEMATODES<br />
Rotylenchulus reniformis Trinidad Singh 1974<br />
VIRUSES<br />
sugarcane mosaic Hawaii, Japan, Taiwan Chen et al. 1989b,<br />
Holm et al. 1937,<br />
Ohtsu & Gomi 1985
184 Biological Control of Weeds: Southeast Asian Prospects<br />
Passiflora foetida<br />
(after Soe jani et a/. 1987)
Map 4.22 Passiflora foetida<br />
4.22 Passiflora foetida<br />
Passiflora foetida<br />
No searches have been made for natural enemies of P. foetida in its area of origin in<br />
South America, where it is not a weed. It is known to be attacked there by the larvae of<br />
some nymphalid (heliconiine) butterflies.<br />
However studies of the related P. tripartita, a serious forest weed in Hawaii,<br />
recorded upwards of 200 species of insects. It may thus be inferred that a similar study<br />
would reveal many insects attacking P. foetida. However, until a relevant study is carried<br />
out, it is not possible to evaluate the prospects for its successful biological control.<br />
I
186 Biological Control of Weeds: Southeast Asian Prospects<br />
4.22 PassifZora foetida L.<br />
Passifloraceae<br />
stinking passionflower, wild passionfruit; love-in-a-mist; ka thok rok (Thailand),<br />
timun padang (Malaysia), buah tikus (Indonesia)<br />
Rating<br />
+++ Msia<br />
11 ++ Brun<br />
+ Myan, Thai, Laos, Viet, Sing, Phil<br />
Indo<br />
Origin<br />
South America. Natural populations have been observed in the coastal mountain ranges<br />
in the State of Parana, Brazil (G.P. Markin pers. comm. 1993).<br />
Distribution<br />
Widespread throughout the tropics and serious in Southeast Asia; also a weed in the<br />
Pacific Region, West Africa and Central America. Introduced to Java a long time ago.<br />
Characteristics<br />
A foetid, woody, annual or perennial vine, 1.5 to 6 m long; stem, cylindrical, densely<br />
hairy; tendrils arise next to leaves on the shaded side of the stem; leaves heart-shaped to<br />
three lobed, alternate, arranged helically, with long-stalked glands and long fine hairs on<br />
margins, producing a disagreeable smell when crushed; flowers white to lilac, bisexual. It<br />
flowers all year round, opening in the morning and closing before noon. The green to<br />
orange or red fruits are enclosed in lacy bracts. A large number of varieties occur<br />
(Wagner et al. 1990).<br />
Importance<br />
I! foetida is a weed of upland rice and other field crops. It occurs in wet areas or those<br />
where there is a pronounced wet season. It is common in plantations, rough pastures,<br />
roadsides and wasteland.<br />
In the Philippines it is sometimes used as a soil cover in coconut plantations to con-<br />
trol Imperata cylindrica grass or erosion. In Papua New Guinea it is planted between<br />
sweet potatoes to suppress Imperata. Young leaves are used in Surinam and Java as a<br />
vegetable. Seeds are flat, black, woody and enclosed in a sweet aromatic pulp (Swarbrick<br />
1981). Young fruit are cyanogenic. Stems and leaves are suspected of poisoning live-<br />
stock.<br />
P. foetida contains alkaloids and at least 10 flavonoids. One of the latter, ermanin, is<br />
a feeding deterrent to larvae of the nymphalid butterfly Dione juno which, in Colombia,<br />
do not attack P. foetida leaves, but eat large amounts of other PassiJlora species.
4.22 Passifora foetida 187<br />
The Passifloraceae contain about 12 genera and 600 species, most of which are ten-<br />
dril climbing vines native to warm regions of the world. The genus PassiJlora contains<br />
some 500 tropical and subtropical species, mostly from Central and South America.<br />
Several have edible fruits and attractive flowers, about 40 species have been cultivated,<br />
but fewer than 6 are fruit crops in the neotropics and only one, P. edulis (and its varieties,<br />
such as the yellow favicarpa), is economically important (Waage et al. 1981). P. ligu-<br />
laris is also cultivated in Malaysia (Ong and Ting 1973). A few species, such as P. foeti-<br />
da and P. lonchocarpa, are extremely foul smelling (Benson et al. 1976). Eleven species,<br />
including P. foetida and P. tripartita (= P. mollissima) (in Hawaii) are recorded as weeds<br />
in different parts of the world (Swarbrick 1981). Both P. foetida and P. tripartita are<br />
closely related taxonomically, whereas, P. edulis belongs to a different subgenus (Waage<br />
et al. 1981) and is the only economic crop at risk from oligophagous insects attacking<br />
P. foetida.<br />
Natural enemies<br />
Upwards of 200 insects are recorded attacking Passifloraceae in Central and South<br />
America. The most notable are heliconiine butterflies of the family Nymphalidae. Their<br />
larvae develop only on plants of the family Passifloraceae, with the single exception of<br />
Eueides procula, which will develop on the Turneraceae (Pemberton 1983, Waage et al.<br />
1981). only 5 of the 65 or so species of heliconiines are recorded as pests of Passiflora<br />
edulis, namely Agraulis vanillae, Dione juno, Dryas julia, Eueides aliphera and E.<br />
isabella, although larvae of a few other species are occasionally found on it (Waage et al.<br />
1981). It is apparent that heliconiine butterflies are well worth investigating for species of<br />
adequate host specificity to P. foetida.<br />
Little is known about the natural enemies of P. foetida (Table 4.22.1) and no<br />
attempts have been made at biological control. The passion vine butterfly Agraulis vanil-<br />
lae, an accidental introduction to Hawaii before 1977, is now widespread there. In addi-<br />
tion to attacking Passifora edulis, its larvae feed on the leaves of P. foetida, P. manicata<br />
and P. suberosa, but they seldom attack banana poka, P. tripartita, which is a serious for-<br />
est weed in Hawaii. P. foetida is widely distributed on Hawaii from sea level up to about<br />
500 m and a rainfall from 750 to 3000 mm. It occurs generally in highly disturbed areas,<br />
where it is a very minor component among other introduced species. On the west side,<br />
the taxon has red fruit and, on the east, green. It has very few natural enemies, with the<br />
exception of Agraulis vanillae. A. vanillae larvae are common but usually in small num-<br />
bers, although occasionally there are outbreaks that completely defoliate the plants (G.P.<br />
Markin pers. comm. 1993). A. vanillae is native to the Americas and ranges from<br />
Argentina up through Mexico to Florida, the Gulf States and California (Beardsley 1980,<br />
Bianchi 1982, 1983, Klots (1951)). In Hawaii it is attacked by a nuclear polyhedral virus<br />
which limits its numbers (G.P. Markin pers. comm. 1993), in California by Phorocera<br />
claripennis (Tachinidae) and in eastern USA by Brachymeria ovata (Nakahara 1977).<br />
The other species of heliconiine recorded as attacking P. foetida is Heliconius hecale,<br />
which is widespread in Central and South America and attacks a large number of
188 Biological Control of Weeds: Southeast Asian Prospects<br />
Passifloraceae (Benson et al. 1976, Waage et al. 1981). On the other hand, larvae of<br />
H. charithonia, H. cydno and H. erato did not develop on R foetida (Waage et al. 1981).<br />
In the Ivory Coast larvae of the pterophorid moth Sphenarches anisodactylus eat the<br />
leaves of R foetida, Lagenaria siceraria and Brillantaisia lamium. Although the moth<br />
also occurs in India and Japan it is not known from P. foetida there, but attacks two eco-<br />
nomic plants, the legumes lablab bean, Lablab purpureus and pigeon pea, Cajanus cajan<br />
(Bigot and Vuattoux 1979). Thus there is some uncertainty whether the host specificity of<br />
the African taxon is the same as that in Asia.<br />
The National Parks and Forest Service authorities in Hawaii have been carrying out<br />
searches for some years in South America for natural enemies of banana poka, R triparti-<br />
ta. Two insects from Colombia have been introduced to Hawaii (Gardner et al. 1992).<br />
One of these was the moth Cyanotricha necyria (Dioptidae), which was released in 1988,<br />
but did not become established (Casaiias-Arango et al. 1990, Julien 1992, Markin and<br />
Nagata 1989, Markin et al. 1989). In host specificity tests C. necyria did not oviposit on<br />
f? foetida, but the larvae could develop on its foliage (Markin and Nagata 1989). The<br />
fungus Colletotrichium gloeosporioides f. sp. clidemiae has been mass produced for lib-<br />
eration (E.E. Trujillo memorandum 1989).<br />
In Hawaii the fungus Fusarium oxysporum f. sp. passiflorue attacks P. foetida,<br />
P. tripartita and P. ligularis, but not R suberosa or the cultivated P. edulis f. jlavicarpa<br />
(Gardner 1 989).<br />
Table 4.22.1 Natural enemies of Passiflora foetida.<br />
Species Location Other hosts References<br />
INSECTS<br />
Hemiptera<br />
APHlDlDAE<br />
Aphis fabae Kenya polyphagous Bakker 1974<br />
Aphis gossypii Ivory Coast polyphagous De Wijs 1974<br />
Aphis spiraecola Ivory Coast De Wijs 1974<br />
Myzus persicae Japan polyphagous Yonaha et al. 1979<br />
Uroleucon compositae Kenya Bakker 1974<br />
Diptera<br />
AGROMYZIDAE<br />
Melanagromyza<br />
PO~YP~Y~~<br />
Tropicomyia theae<br />
Lepidoptera<br />
NOCTUIDAE<br />
Helicoverpa zea<br />
(= H. obsoleta)<br />
. Heliothis virescens<br />
NYMPHALIDAE<br />
Agraulis vanillae<br />
f<br />
Australia polyphagous, including Kleinschmidt 1960, 1970<br />
Passijlora spp.<br />
Papua New polyphagous Spencer I990<br />
Guinea<br />
Sumatra polyphagous Den Doop 19 18<br />
Venezuela polyphagous Venturi 1960<br />
Central restricted to some Anon 1977, Beardsley 1980,<br />
America, Passijlora spp. Bianchi 1982, 1983,<br />
Hawaii Waage et al. 198 1<br />
(continued on next page)
FUNGI<br />
4.22 Passifloi-a foetida 189<br />
Species Location Other hosts References<br />
Heliconius hecale<br />
PTEROPHORIDAE<br />
Sphenarches<br />
anisodactylus<br />
Alternaria passiflorae<br />
Alternaria tenuis<br />
Colletotrichium<br />
gloeosporioides<br />
Fusarium oxysporum<br />
f.sp. passiflorae<br />
Haplosporella<br />
pass$oridia<br />
Hemphyllium sp.<br />
NEMATODE<br />
Meloidogyne incognita<br />
VIRUS<br />
cucumber mosaic<br />
passionfruit chlorotic<br />
spot<br />
passionfruit moiaic<br />
passionfruit ringspot<br />
potyvirus<br />
passionfruit woodiness<br />
potyvirus<br />
widespread in restricted to some<br />
Central & South Pass$ora spp.<br />
America<br />
Ivory Coast see discussion<br />
Hawaii<br />
Hawaii<br />
India<br />
Hawaii<br />
India<br />
Hawaii<br />
Australia<br />
Japan<br />
Papua New<br />
Guinea<br />
Hawaii,<br />
Malaysia<br />
Ivory Coast<br />
Australia,<br />
Kenya<br />
Waage et al. 198 1<br />
Bigot & Vuattoux 1979<br />
Raabe 1965<br />
Raabe 1965<br />
Mallikarjunaiah & Rao 1972<br />
Gardner 1989<br />
Pande 1980<br />
Raabe 1965<br />
Sauer & Alexander 1979<br />
Yonaha et al. 1979<br />
Van Velsen 196 1<br />
Ong & Ting 1973,<br />
Raabe 1965<br />
Brunt et al. 1990,<br />
De Wijs 1974<br />
Anon 1976, Bakker 1974,<br />
Brunt et al. 1990,<br />
Leggat & Teakle 1975
190 Biological Control of Weeds: Southeast Asian Prospects<br />
Pennisetum polystachion<br />
(after Holm et a/. 1977)
Map 4.23 Pennisetum polystachion<br />
4.23 Pennisetum polystachion 191<br />
Pennisetum polystachion<br />
This erect, tufted, non-stoloniferous grass, originated in Tropical Africa from where it<br />
has spread throughout Asia and Southeast Asia to the Pacific.<br />
Almost nothing is known of the natural enemies of P. polystachion or closely<br />
related species. It is not possible to evaluate the prospects for its biological control with-<br />
out a search for natural enemies in its region of origin.
192 Biological Control of Weeds: Southeast Asian Prospects<br />
4.23 Pennisetum polystachion (L.) Schultes<br />
(= Pennisetum setosum)<br />
Poaceae<br />
mission grass; feather Pennisetum; yaa khaehyon chop (Thailand), rumput<br />
gajah, rumput berus, rumput kuning, ekor kucing (Malaysia) rumput jurig<br />
(Indonesia)<br />
There are differences in opinion over the spelling of the specific name, polystachion or<br />
polystachyon, with the former being used here. In Africa, there are three subspecies<br />
P. polystachion polystachion, P. p. setosum (sometimes regarded as a true species) and<br />
P. p. atrichum. There is some evidence of crossing between the varieties of P. polysta-<br />
chion and the related Pennisetum hordeoides and the production of populations with dif-<br />
ferent chromosome numbers (Brunken 1979).<br />
Rating<br />
+++ Msia<br />
11 ++ Indo, Phil<br />
+ Thai, Laos, Camb, Viet<br />
Origin<br />
Tropical Africa.<br />
Distribution<br />
P. polystachion is widespread in the tropics of Africa and Asia, but also occurs in north-<br />
ern Australia and the Pacific. It rarely extends beyond 23"N or 23"s. In Africa it occurs<br />
mainly in the savanna and open areas in the forest zone of West Africa from Senegal to<br />
Cameroun and then south and east to Mozambique and Kenya (Brunken 1979, Kativu<br />
and Mithen 1988).<br />
Characteristics<br />
P. polystachion is an erect, tufted annual or perennial grass, with fibrous roots, but no<br />
stolons. Its leaves are narrow and 5 to 45 cm long. Its flowering stems are sometimes<br />
branched, 50 to 300 cm tall, ending in a cylindrical yellow-brown flowering spike, 5 to<br />
25 cm long, bearing densely hairy, unequal bristles of two lengths, the longer 2 to 5 cm<br />
and the shorter lcm.<br />
Importance<br />
P. polystachion grows on dry lateritic soils and is often present along roadsides, in waste-<br />
lands and in upland crops. Propagation is by seeds, but regrowth can occur from dormant<br />
buds located at the base of the stems and from aerial nodes.<br />
It becomes dominant in upland tropical hillsides and croplands after forests have
4.23 Pennisetum polystachion 193<br />
been cleared, or when shifting cultivation or subsistence agriculture have been practised<br />
(Holm et al. 1977). Because of the rapid germination of its wind-dispersed seeds and its<br />
aggressive and highly competitive growth, it rapidly takes over wastelands. Since a sin-<br />
gle cultivation rarely kills enough of the weed to provide control, it often impedes further<br />
use of areas for crops.<br />
In Indonesia it was first observed in 1972 (Titrosoedirdjo 1990). It is now an impor-<br />
tant weed of rubber and occasionally a problem in upland rice.<br />
P. polystachion is thought to have reached Malaysia via Thailand as recently as the<br />
early 1980s, infesting at least 10 km2 of roadsides and is now widely distributed, occur-<br />
ring up to an altitude of 900 m (Baki et al. 1990). It is now a major weed in rubber, oil<br />
palm, sugarcane, orchards, vegetables and upland rice (Titrosoedirdjo 1990).<br />
In the Philippines P. polystachion is able to compete effectively even with blady<br />
grass Imperata cylindrica in grassy fields in Central Luzon and in rubber plantations in<br />
west Java (Titrosoedirdjo 1990).<br />
P. polystachion is a good fodder grass when young and makes excellent hay.<br />
Natural enemies<br />
Very few natural enemies of P. polystachion appear to have been recorded in the litera-<br />
ture (Table 4.23.1). The only species of possible relevance are several gall midges from<br />
Africa, but very little is recorded of their biology. Three species of gall midge have been<br />
reared from P. polystachion in the Gold Coast. One is similar to the pestiferous sorghum<br />
midge Contarinia sorghicola (but may be different), the second belongs to the Trifila<br />
group and the third belongs to the Lasiopterariae (Barnes 1954a,b, Geering 1953). Three<br />
species of gall midge have been described from ears of Pennisetum in southern India,<br />
Cecidomyia penniseti (from P. glaucum = P. typhoideum), Geromyia (= Itonida) penniseti<br />
(from P. cenchroides) and Geromyia (= Itonida) seminis and an unidentified species from<br />
the stems. Of these, G. penniseti may be predaceous (Barnes 1954b, Felt 1920, 1921).<br />
In Madagascar there is a gall midge (?Cecidomyia sp.) whose larvae live in the<br />
inflorescence of Pennisetum (no species given) and in Sudan a gall midge, possibly<br />
Geromyia seminis, has been reared from the ears of Pennisetum (no species given)<br />
(Barnes 1954b).<br />
The larvae of the Brazilian satyrid butterfly Eryphanis polyxena were bred in the<br />
laboratory on P. polystachion (= P. setosum) (Dias 1979), but damage is not reported<br />
from the field.<br />
Comment<br />
There is little doubt that a range of natural enemies attacking f? polystachion would be<br />
found if a search were made in Tropical Africa.<br />
Several other weedy species of Pennisetum also originated in Africa, in addition to<br />
three or more species that have at least some desirable attributes. Perhaps the best known is<br />
kikuyu grass, P. clandestinum, which is a valuable fodder during the warmer months,<br />
although it is a weed in some situations and its nitrate levels can be toxic to grazing animals.<br />
?? glaucum (= P. americanum), pearl millet, is used as food in some areas. ?? purpureum,
194<br />
Biological Control of Weeds: Southeast Asian Prospects<br />
elephant or napier grass, can grow to 3 m, is valuable for fodder when young and can be<br />
used as fuel when old. However, it can also be a weed, as in rubber in Malaysia. Little<br />
has been recorded of the natural enemies attacking these species of Pennisetum. The<br />
most interesting are three cecidomyiid gall midges in India, Geromyia penniseti, G. sem-<br />
inis and Mycodiplosis indica from Pennisetum glaucum (= P. typhoideum) and Geromyia<br />
seminis from Pennisetum cenchroides (Barnes 193 1, Felt 1920).<br />
Table 4.23.1 Natural enemies of Pennisetum polystachion.<br />
Species<br />
INSECTS<br />
Diptera<br />
CEClDOMYllDAE<br />
Cecidomyia penniseti<br />
Contarinia ?sorghicola<br />
Contarinia sp. 1<br />
Contarinia sp. 2<br />
Contarinia sp. 3<br />
Contarinia sp. 4<br />
Geromyia (= Itonida) penniseti<br />
Geromyia (= Itonida) seminis<br />
Lepidoptera<br />
HESPERllDAE<br />
Parnara bada bada<br />
NYMPHALIDAE<br />
Eryphanis polyxena<br />
PYRALlDAE<br />
Cnaphalocrosis medinalis<br />
FUNGI<br />
Bipolaris papendor-i<br />
Gloeocercospora sp.<br />
Helminthosporiurn rostratum<br />
Phakospora apoda<br />
Puccinia chaetochloae<br />
Puccinia substrata<br />
Pyricularia oryule<br />
Spacelotheca penniseti<br />
Country<br />
India<br />
Gold Coast<br />
Gold Coast<br />
Gold Coast<br />
Madagascar<br />
India<br />
India<br />
India. Sudan<br />
Malaysia<br />
Brazil<br />
Malaysia<br />
Brazil<br />
References<br />
Barnes 1954b<br />
Barnes 1954b<br />
Bames 1954b<br />
Barnes 1954b<br />
Barnes 1954b<br />
Bames 1954b<br />
Barnes 1954b<br />
Barnes 1954b<br />
C.L. Tan pers. comm. 1993<br />
Dias 1979<br />
C.L. Tan pers. comm. 1993<br />
H.C. Evans pers. comm. I992<br />
H.C. Evans pers. comm. 1992<br />
Thite & Chavan 1977<br />
H.C. Evans pers. comm. 1992<br />
H.C. Evans pers. comm. 1992<br />
H.C. Evans pers. comm. 1992<br />
Prabhu et al. 1992<br />
H.C. Evans pers. comm. I992
196 Biological Control of Weeds: Southeast Asian Prospects<br />
Pistia stratiotes<br />
(after Holm et a/. 1977)
Map 4.24 Pistia stratiotes<br />
4.24 Pistia stratiotes<br />
Pista stratiotes<br />
Water lettuce is a widespread, floating water weed, which probably originated in South<br />
America.<br />
The host specific South American weevil, Neohydronomus aflnis, has been<br />
established readily in six countries and, in all, has produced substantial to excellent con-<br />
trol. The moth, Samea multiplicalis, which attacks P. stratiotes and Salvinia spp., has<br />
been established in Australia but its impact has not been evaluated.<br />
In Thailand, classical biological control has not been attempted, but mass rearing<br />
and release of the native noctuid moth Spodoptera pectinicornis has replaced the use of<br />
herbicides.<br />
The prospects are excellent for classical biological control of P. stratiotes in<br />
countries where it is still regarded as an important weed.
198 Biological Control of Weeds: Southeast Asian Prospects<br />
4.24 Pistia stratiotes L.<br />
Araceae<br />
water lettuce; chak thom (Cambodia), apoe apoe, apon apon (Indonesia),<br />
kiambang besar (Malaysia), chok, jawg (Thailand), beo cai (Vietnam)<br />
Rating<br />
++ Thai, Laos, Camb, Viet, Phil<br />
14 + Myan, Msia, Brun, Indo<br />
Sing<br />
Origin<br />
The origin of Pistia stratiotes is unknown, although the number of host specific insects<br />
present there (Table 4.24.1) suggests South America. However, there is also a host spe-<br />
cific noctuid moth that ranges from India to Papua New Guinea. Dray and Center (1992)<br />
examine the various theories concerning the area of origin of water lettuce.<br />
Distribution<br />
Water lettuce occurs very widely as a troublesome water weed between the tropics of<br />
Capricorn and Cancer, particularly in Africa, Asia, Southeast Asia and the Caribbean.<br />
Pliny refers to its use in Egypt in AD77 (DeLoach et al. 1979, Holm et al. 1977). It is<br />
absent from a number of Pacific countries although recorded as a weed in Papua New<br />
Guinea, Solomon Is, Guam, New Caledonia and Hawaii.<br />
Characteristics<br />
Pistia stratiotes is a free-floating, perennial monocotyledon, with a tuft of fibrous feath-<br />
ery roots up to 1 m long. Numerous secondary roots may be up to 4 cm in length. Stolons<br />
up to 60 cm are produced from the base of the plant and develop into new plants. Leaves<br />
are pale green, upright, 2.5 to 15 cm long, broad at the top and tapered at the base. They<br />
are prominently veined beneath and form a rosette. They are spongy in texture and bear<br />
numerous fine, water-repelling hairs on both sides. The flowers are bisexual, inconspicu-<br />
ous, green, surrounded by tubular bracts and arise from the centre of the rosette. It is said<br />
that water lettuce does not flower in Thailand but a small number of flowers have been<br />
observed (B. Napompeth pers. comm. 1993). It flowers in the Philippines, Australia,<br />
Africa and USA. The fruit is berry-like and green and contains 4 to 12 small brown seeds<br />
which can float on the water for up to 2 days. There are as many as 9 varieties of water<br />
lettuce (Neal 1965).<br />
Importance<br />
The free-floating plants are found in reservoirs, ponds, in marshes along the edges of<br />
large tropical lakes and in slow-moving or stagnant waters. They multiply rapidly and
4.24 Pistia stratiotes 199<br />
can block streams, interfere with fisheries and hydroelectric generating plant and bank up<br />
at dams, bridges and culverts, leading to increased flooding problems. Despite earlier<br />
claims (Holm et al. 1977), unlike water hyacinth, water lettuce does not increase water<br />
loss through evapotranspiration (Lallana et al. 1987). Together with water hyacinth it is a<br />
common and important component of the dense aggregations of free-floating vegetation,<br />
known as sudds. It grows best at pH 4, whereas water hyacinth produces greatest dry<br />
weight at pH 7 (Holm et al. 1977).<br />
Water lettuce plants act as a substrate for sandfly larvae (Ceratopogonidae) and lar-<br />
vae and pupae of the disease-transmitting mosquito genus Mansonia obtain their oxygen<br />
by attaching to Pistia roots. P. stratiotes is an important weed of irrigated rice, floating<br />
into paddy crops, taking root in the soil and competing much like other weeds.<br />
On the other hand, it has been used as human food in India during famines and is<br />
still fed to pigs and ducks. It is said to have some medicinal value as a cure for skin dis-<br />
eases and dysentery, as a laxative, to treat asthma and, its ash rubbed into the scalp, as a<br />
treatment for ringworm.<br />
Natural enemies<br />
AUSTRALIA<br />
Although water lettuce was first recorded in the Northern Territory only in 1946 it was<br />
already an important weed in some locations in Queensland before the introduction of<br />
Neohydronomus afJinis, although plants in the Northern Territory were rarely thrifty.<br />
Heavy damage was observed there by larvae of the moth Parapoynx (= Nymphula) tene-<br />
bralis, which lays it eggs on the leaves. Newly emerged larvae excise a portion of leaf to<br />
make a protective case in which they shelter while feeding and moving around the plant.<br />
These larvae also attack Salvinia molesta, as do larvae of the related Parapoynx<br />
(= Nymphula) turbata (Gillett et al. 1988). In Thailand this species attacks water lettuce<br />
(Napompeth 1982), so it presumably has the same habit in the Northern Territory. A bug,<br />
Nisia nervosa (= N. atrovenosa) feeds on water lettuce as it does in India (Gillett et al.<br />
1988, Joy 1978).<br />
CENTRAL AND SOUTH AMERICA<br />
Bennett (1975) listed 12 species of phytophagous insects that had been reared from Pistia<br />
stratiotes and Cordo et al. (1981) added one more. Particularly notable (Table 4.24.1) is<br />
the group of South American weevils which are generally confined to water lettuce,<br />
although adults of several may produce minor attack on some of the nearby aquatic<br />
plants whose stems and leaves nevertheless would be too small to support larval develop-<br />
ment. Preliminary host range studies suggest that several may be specific enough to be<br />
employed for biological control, although adequate information is available only for<br />
Neohydronomus afinis. This has been successfully established in several countries (see<br />
later). Larvae of the small pyralid moth Samea multiplicalis, which occurs from<br />
Argentina to the southeastern United States, feed on the growing buds of water lettuce<br />
and sporadically cause very heavy damage and dieback of the plants (Cordo et al. 1978,<br />
1981, DeLoach et al. 1976, 1979).
200 Biological Control of Weeds: Southeast Asian Prospects<br />
Table 4.24.1 Natural enemies of Pistia stratiotes.<br />
Species Location References<br />
INSECTS<br />
Hemiptera<br />
APHlDlDAE<br />
unnamed<br />
Rhopalosiphum nymphaeae<br />
COCCIDAE<br />
Planococcus citri<br />
LYGAEIDAE<br />
Valtissius sp.<br />
MEENOPLllDAE<br />
Nisia nervosa<br />
(= N. atrovenosa)<br />
Orthoptera<br />
ACRlDlDAE<br />
Paulinia acuminata<br />
TETRlGlDAE<br />
Criotettrix sp.<br />
Coleoptera<br />
CURCULIONIDAE<br />
Argentinorhynchus bennetti<br />
Argentinorhynchus breyeri<br />
Argentinorhynchus bruchi<br />
Argentinorhynchus minimus<br />
Argentinorhynchus squamosus<br />
Neohydronomus aflnis<br />
Neohydronomus elegans<br />
Neohydronomus pulchellus<br />
Ochetinu bruchi<br />
Pistiacola cretatus<br />
Pistiacola fasciatus<br />
Pistiacola sp. nr nigrirostris<br />
Lepidoptera<br />
ARCTllDAE<br />
Spilosoma virginica<br />
NOCTUIDAE<br />
Erastroides curvifascia<br />
Proxenus hennia<br />
,Spodoptera pectinicornis<br />
Thailand<br />
Florida<br />
Nigeria, Trinidad<br />
Argentina<br />
Australia. India<br />
South America<br />
Thailand<br />
Mexico, Venezuela<br />
Argentina, Paraguay<br />
Argentina, Paraguay<br />
Venezuela<br />
Argentina, Paraguay<br />
South America<br />
Honduras, Cuba<br />
Trinidad to Argentina, Cuba<br />
Argentina<br />
Argentina, Brazil<br />
Central and South America<br />
Argentina<br />
Florida<br />
India<br />
Indonesia<br />
India, Bangladesh, Thailand<br />
Napompeth 1990a<br />
Ballou et al. 1986, Bennett<br />
1975, Joy 1978<br />
Bennett 1975<br />
Bennett 1975<br />
Gillett et al. 1988, Joy 1978<br />
Bennett 1966, 1975<br />
Napompeth 1990a<br />
O'Brien & Wibmer 1989a,b<br />
O'Brien & Wibmer 1989a,b<br />
Cordo et al. 1978<br />
O'Brien & Wibmer 1989a,b<br />
Cordo et al. 1978<br />
DeLoach et al. 1976<br />
O'Brien & Wibmer 1989c<br />
Bennett et al. 1975,<br />
O'Brien & Wibmer 1989c<br />
Cordo et al. 198 1<br />
Cordo et al. 198 1<br />
Wibmer & O'Brien 1989<br />
Cordo et al. 198 1<br />
Thompson and Habeck 1988<br />
Chaudhuri & Janaki Ram<br />
1975<br />
Mangoendihardjo & Nasroh<br />
1976<br />
Alam et al. 1980, George<br />
1963, Napompeth 1990a,<br />
Sankaran 1974, Sankaran &<br />
Rao 1972, Sankaran et al.<br />
1964<br />
(continued on next page)
MITES<br />
FUNGI<br />
4.24 Pistia stratiotes 201<br />
Species Location References<br />
PYRALlDAE<br />
Argyractis subornata<br />
Elophila responsalis<br />
Parapoynx (= Nymphula)<br />
diminutalis<br />
Parapoynx (= Nymphula)<br />
tenebralis<br />
Parapoynx (= Nymphula)<br />
turbata<br />
Petrophila drumalis<br />
Samea multiplicalis<br />
Synclita obliteralis<br />
Hydrozetes subornata<br />
Cercospora canescens<br />
Cercospora sp.<br />
Phyllosticta stratiotes<br />
Sclerotium rolfsii<br />
Brazil<br />
India, Indonesia<br />
Thailand<br />
Australia<br />
Australia, Thailand<br />
Florida<br />
southern USA, Trinidad,<br />
northern South America<br />
Florida<br />
Australia<br />
Australia<br />
India<br />
India<br />
India<br />
Fomo 1983<br />
Handayani & Syed 1976,<br />
Mangoendihardjo et al. 1977,<br />
Sankaran & Rao 1972,<br />
Subagyo1975<br />
Napompeth 1990a,<br />
Suasa-Ard 1976<br />
Gillett et al. 1988<br />
Gillett et al. 1988,<br />
Napompeth 1990a,<br />
Suasa-Ard 1976<br />
Dray et al. 1988<br />
Bennett 1966, 1975,<br />
Bennett et al. 1975,<br />
Dray et al. 1988<br />
Dray et al. 1988<br />
Gillett et al. 1988<br />
Gillett et al. 1988<br />
Bennett 1975,<br />
Nag Raj and Ponappa 1966<br />
Bennett 1975<br />
Bennett 1975<br />
UNITED STATES<br />
Dray et al. (1988) recorded larvae of three species of moth, a mealy bug, a leafhopper<br />
and an aphid on water lettuce which has been present in Florida for at least 200 years<br />
(Thompson and Habeck 1988). Only one of these insects, a root feeding moth, was con-<br />
sidered to be possibly host specific. This was later identified as the pyralid moth<br />
Petrophila drumalis: the two other moths were Samea multiplicalis and Synclita obliter-<br />
alis (Dray et al. 1989). The aphid was probably Rhopalosiphum nymphaeae, a well<br />
known transmitter of a number of economically important viruses. It has been recorded<br />
to cause dieback of water lettuce in Nigeria (Pettett and Pettett 1970). This aphid was<br />
reported to be widespread on water lettuce in Florida (Ballou et al. 1986). An aphid, pos-<br />
sibly the same species, transmitted a virus that caused widespread dieback of Z? stratiotes<br />
on Lake Volta in Ghana (Okali and Hall 1974), although serious dieback has not been<br />
reported in Florida. The non-specific larvae of the arctiid moth Spilosoma virginica was<br />
also common on water lettuce in Florida (Thompson and Habeck 1988).
202 Biological Control of Weeds: Southeast Asian Prospects<br />
INDIA<br />
Larvae of the moth Spodoptera pectinicornis cause extensive damage to Pistia. On aver-<br />
age, a single larva can consume the leaves of two Pistia plants during its developmental<br />
period of 15 to 20 days. Some 100 larvae developing from an average egg mass destroy<br />
all Pistia leaves within an area of 1 m2 and, during peak abundance in the field, the num-<br />
ber of larvae per m2 of Pistia surface was always higher than this (George 1963,<br />
Sankaran and Ramaseshiah 1974). The bug Nisia nervosa successfully completes its life<br />
cycle on Pistia, but is reported as a minor pest of rice (Joy 1978).<br />
INDONESIA<br />
In Java and Sulawesi, water lettuce is attacked by larvae of the noctuid moth Proxenus<br />
hennia which appears to be specific (Mangoendihardjo and Nasroh 1976). Other species<br />
found attacking it were Elophila (= Nymphula) responsalis, Spodoptera mauritia, an<br />
aphid and a cicadellid (Zygina sp.) (Mangoendihardjo and Syed 1974, Mangoendihardjo<br />
et al. 1976, 1977, Syed et al. 1977).<br />
THAILAND<br />
Water lettuce is attacked by several insects (Table 4.24.1), of which only the pygmy<br />
grasshopper Criotettrix sp. and the native water lettuce moth Spodoptera pectinicornis<br />
are capable of inflicting serious damage. In certain areas where the density of Criotettrix<br />
was as high as 100 per m2 considerable suppression of the weed occurred. Both adults<br />
and nymphs were able to walk on the surface of the water and were observed to attack<br />
also the water fern, Salvinia cucullata (Napompeth 1982). The extensive damage that can<br />
be caused by Spodoptera pectinicornis is discussed later.<br />
Attempts at biological control<br />
AUSTRALIA<br />
The first attempt to bring about classical biological control of Pistia stratiotes was the lib-<br />
eration of adults and larvae of Neohydronomus afinis in 1982 near Brisbane. Within two<br />
months of release, plants were rotting and sinking and, by eight months, about one third<br />
of the plants in a dam were chlorotic and some had been destroyed. Severely damaged<br />
plants produced short stolons terminating in small plantlets before sinking and dying, but<br />
these plantlets failed to grow to the size of their parents before, in turn, becoming severe-<br />
ly damaged, producing plantlets and then sinking. Continued weevil attack led initially to<br />
an increase in the number of plants, but a decrease in their size and dry weight. Before<br />
long, few water lettuce plants remained (Harley et al. 1984). The moth Samea multipli-<br />
calis was liberated in Australia in 1981, primarily against Salvinia molesta, on which it<br />
became established. However, within four years, its effectiveness was restricted by proto-<br />
zoan disease and three hymenopterous parasitoids (Thomas and Room 1986). It presum-<br />
ably attacks Pistia stratiotes also, although there seems to be only one observation of it<br />
doing so. This was at Townsville (D.P.A. Sands pers. comm. 1993).<br />
PAPUA NEW GUINEA<br />
The moth Spodoptera pectinicornis attacks water lettuce, but is unable to prevent its<br />
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.<br />
4.24 Pistia stratiotes 203<br />
Species Where From When Result References<br />
Coleoptera<br />
CURCULIONIDAE<br />
Neohydronornus a#nis Australia Brazil 1982 + Harley et al. 1984, 1990<br />
Botswana Brazil via 1988 + Chikwenhere & Forno 1991<br />
Australia LW. Fomo pers. comm. 1993<br />
Papua New Brazil via 1985 + Chikwenhere & Forno 199 1,<br />
Guinea Australia Harley et al. 1990,<br />
Laup 1987b<br />
South Africa Brazil via 1985 + Cilliers 1987, 1989b<br />
Australia<br />
United States Brazil via 1987 + Center et al. 1989,<br />
of America Australia Thompson & Habeck 1988,<br />
Dray et al. 1990<br />
Zambia Zimbabwe about + P. Room pers. comm. 1993<br />
1990<br />
Zimbabwe Brazil via 1988 + Chikwenhere & Fomo 1991<br />
Australia<br />
Lepidoptera<br />
NOCTUIDAE<br />
Spodoptera pectinicornis Florida Thailand 1990 ? Center et al. 1989,<br />
Julien 1992<br />
Napompeth 1990a<br />
PYRALl DAE<br />
Sarnea rnultiplicalis Australia Brazil 198 1 + Forno 1987, Room et al. 1984<br />
Eichhornia. Neohydronomus afinis was successfully established in the Sepik River sys-<br />
tem in 1985, but its impact is yet to be recorded (Laup 1987b).<br />
THAILAND<br />
Although no introductions of biological control agents for Pistia stratiotes have been<br />
made in Thailand, the mass rearing and release of the native noctuid moth Spodoptera<br />
pectinicornis has replaced the use of herbicides for this weed. Under laboratory condi-<br />
tions mixed instar larvae at the rate of 300 or more per m2 gave as fast and effective con-<br />
trol as any herbicide. In the field a substantial initial release of larvae, followed by one or<br />
two additional releases at two-week intervals has led to complete control within 6 to 10<br />
weeks. Thus, a 4.5 km2 infestation of water lettuce was controlled in 6 weeks at Sri<br />
Nakarint Dam in 1978 and a 10 km2 infestation in 1982 (Napompeth 1982). S. pectini-<br />
cornis occurred throughout the year and in all infestations of Pistia (Suasa-Ard and<br />
Napompeth 1982).<br />
UNITED STATES<br />
Neohydronomus afinis was released in Florida in 1987, became established readily, mul-<br />
tiplied rapidly and soon spread from the release sites to cause considerable damage to<br />
water lettuce (Dray et al. 1990). In one release site the Pistia population was reduced<br />
from 50 to less than 5 acres in 2 years and, in another, a 10 acre site was virtually cleared
204 Biological Control of Weeds: Southeast Asian Prospects<br />
in 3 years. However, in a third site, little effect was noted. It was postulated that this<br />
might be due to the presence of a different genetic strain of P. stratiotes, which had a far<br />
greater seed production than that at the other two sites (Dray and Center 1992).<br />
Spodoptera pectinicornis has also been established in Florida (Center et al. 1989;<br />
Napompeth 1 990a).<br />
BOTSWANA<br />
N. afinis was released on the Linyanti R at the Selinda spillway in 1988. Excellent con-<br />
trol was achieved within 12 months (I.W. Forno pers. comm. 1993).<br />
SOUTH AFRICA<br />
Neohydronomus afinis was released in Kruger National Park and a water lettuce infesta-<br />
tion in a motionless water body was completely controlled within 10 months (Cilliers<br />
1987, 1989a,b). The weevil has been less successful on fast-flowing rivers where plants<br />
infested with weevil larvae are continually washed down stream and replaced by unin-<br />
fested plants from higher up. However, even under these circumstances, up to 90% of<br />
plants showed signs of feeding damage (Cilliers 1991 b).<br />
ZAMBIA<br />
N. afinis was already established by natural spread at Kafubu Lake when N. afinis from<br />
Zimbabwe was liberated about 1990 and by 1992 there were only scattered plants of<br />
Pistia but no mats (P.M. Room pers. comm. 1993).<br />
ZIMBABWE<br />
Neohydronomus afinis was released in 1988, was well established in 4 months and, with-<br />
in a year, water lettuce was no longer a problem in the Manyame River (Chikwenhere<br />
and Forno 1991).<br />
Major natural enemies<br />
Argentinorhynchus bmchi Coleoptera: Curculionidae<br />
This yellow spotted weevil (4.7 mm long) is known from Argentina, Bolivia, Paraguay<br />
and Uruguay (O'Brien and Wibmer 1989a,b). Although it is rare, it has the potential to<br />
cause heavy damage to water lettuce. Under laboratory conditions adults ate 1 cm2 of<br />
leaf surface per day, producing some 10 oval holes all the way through the leaf. Adults<br />
feed mostly by night and generally on medium-aged leaves. Field collected females laid<br />
on average 1575 eggs among the dense hairs on the leaf surface. Eggs hatch in 7.6 days.<br />
First instar larvae enter the leaf and feed on the spongy leaf tissue and second and third<br />
instars in the crown. Fourth instar larvae could not be reared: in the laboratory they left<br />
the plant and drowned. Adults fed and oviposited only on water lettuce and, except for<br />
slight feeding on Spirodela, of the 26 plant species tested, larvae only developed on water<br />
lettuce. In the laboratory 6 larvae per plant killed water lettuce within a month. It was<br />
suggested that egg predation may account for the rarity of A. bruchi; also that the diffi-<br />
culty experienced in rearing the fourth instar larvae may indicate that special conditions<br />
are required, lack of which may reduce survival (Cordo et al. 1978).
4.24 Pistia stratiotes 205<br />
Neohydronomus aflnis Coleoptera: Curculionidae<br />
This mottled, brown-grey weevil was earlier confused in the literature with the closely<br />
related N. pulchellus. It occurs naturally in Argentina, Brazil, Colombia, Paraguay, Peru,<br />
Uruguay and Venezuela (O'Brien and Wibmer 1989~). Adults (males 1.8 mm, females<br />
2.1 mm long) feed on the leaves of Pistia stratiotes and mine the tissues: they do not<br />
appear to attack the crown or roots. Females lay about 1 egg per day beneath the leaf epi-<br />
dermis, usually on the upper surface near the margin. Eggs hatch after 3 to 4 days and<br />
larvae tunnel through the leaf tissues to complete development in 1 1 to 14 days. Pupation<br />
occurs in small pockets in the leaf tissues and adults emerge after about 4 days. The peri-<br />
od from egg to adult varies from 4 to 6 weeks, but there are only 3 generations a year in<br />
Argentina (December, February to March and June). Overwintering probably occurs in<br />
the adult stage (DeLoach et al. 1976).<br />
N. afJinis is very destructive under laboratory conditions. Maximum damage<br />
occurred in midsummer in Argentina, when peak populations of 200 to 600 per m2 pro-<br />
duced 1.6 feeding spots per cm2 of leaf surface. Adult N. afinis are occasionally para-<br />
sitised by nematodes in Argentina (DeLoach et al. 1976).<br />
N. affinis is highly specific to water lettuce, as shown by tests in Zimbabwe<br />
(Chikwenhere and Forno 1991), South Africa (Cilliers 1989b), Florida (DeLoach et al.<br />
1976, Dray et al. 1990, Thompson and Habeck 1988, 1989) and Australia (Harley et al.<br />
1990); and also by absence of reported damage to economic plants in any of the countries<br />
to which it has been introduced.<br />
Pistiacola cretatus Coleoptera: Curculionidae<br />
This brown 2.3mm long weevil, earlier known as Onychylis cretatus, occurs in Argentina<br />
and Brazil (Wibmer and O'Brien 1989). Adults feed mainly on the upper surface of the<br />
leaves of P. stratiotes and oviposit into the leaf tissue. The slender larvae tunnel into the<br />
denser tissues of the basal third of the leaf and also into the crown. Pupation occurs with-<br />
in the spongy part of the leaf. In the field, adult P. cretatus were found only on water let-<br />
tuce (Cordo et al. 1981).<br />
Samea multiplicalis Lepidoptera: Pyralidae<br />
This brown moth with dark markings and a wingspan of about 17 mm occurs from<br />
Florida to Argentina.<br />
Up to 290 eggs are laid per female, mainly on the upper surface of the leaves. These<br />
hatch after 4 days and the larvae construct a silken canopy under which they feed, or they<br />
may tunnel into the leaves to feed on the spongy tissues: they also eat the buds. After 5 or<br />
6 instars in the course of 16 days, they pupate in silken cocoons, to emerge as adults 5<br />
days later. Adults live up to 7 days (DeLoach et al. 1979, Knopf and Habeck 1976, Sands<br />
and Kassulke 1984). S. multiplicalis has three main hosts in Florida, Pistia stratiotes,<br />
Azolla caroliniana and Salvinia rotundifolia and it may occasionally attack Eichhornia<br />
crassipes. Oviposition is highest on P. stratiotes. Although medium to large larvae fed on<br />
a number of plants under laboratory conditions, S. multiplicalis has never been reported
06 Biological Control of Weeds: Southeast Asian Prospects<br />
as a pest of cultivated plants in Argentina or Brazil (DeLoach et al. 1979). It passed strict<br />
host specificity tests in Australia, larvae completing development on l? stratiotes, Azolla<br />
pinnata and Salvinia molesta. Larvae that had fed first on S. molesta were unable to com-<br />
plete their development on water lettuce, although they produced minor leaf scars. S.<br />
multiplicalis was released in Australia, but primarily against Salvinia molesta (Sands and<br />
Kassulke 1984).<br />
Samea multiplicalis has 3 generations a year in the field in Argentina, with popula-<br />
tion peaks in December, February and May, when populations reach a maximum of 5 lar-<br />
vae per plant. In laboratory tests females laid 99.3% of their eggs on P. stratiotes. Larvae<br />
caused heavy, but sporadic, damage to water lettuce in the field. However, in most years,<br />
populations were held at low levels, apparently by parasitoids (Apanteles sp. and<br />
Podogaster sp.) (DeLoach et al. 1979). In Florida 52% parasitisation was recorded,<br />
42.7% by three species of Hymenoptera (Agathis sp., Apsilops sp. and Temelucha ferrug-<br />
inae) and 9.3% by a tachinid fly (Lixophaga sp.) (Knopf and Habeck 1976). Nosema sp.<br />
was detected in Australia in some larvae from Brazil and the culture was freed of these<br />
before release (Sands and Kassulke 1984).<br />
Spodoptera pectinicornis Lepidoptera: Noctuidae<br />
This moth ranges over an extensive area from India through Sri Lanka, Thailand and<br />
Indonesia to Papua New Guinea. Eggs are laid in masses of 70 to 120 on the undersur-<br />
face of the Pistia leaf near its edge. They hatch in 40 to 60 hours to produce pale green<br />
larvae that burrow in the leaf parallel to the longitudinal veins. After some 20 days the<br />
1.5 to 2 cm long larvae pupate, to emerge two to three days later as small silvery brown<br />
moths about lcm long (George 1963). In Thailand the period from egg to adult averaged<br />
30 days and females laid an average of 666 eggs. Host specificity tests showed that it<br />
would develop only on Pistia stratiotes (Suasa-Ard 1976). Napompeth (1990a) reports<br />
that it is relatively simple to mass rear in the laboratory and to distribute in the field.<br />
Details are available of its rearing and ecology in Thailand (Suasa-Ard 1976, Suasa-Ard<br />
and Napompeth 1978). It was mass reared and released in Florida after tests showed that<br />
it was sufficiently host specific (Center et al. 1989, Napompeth 1990a).<br />
Comment<br />
Pistia stratiotes is seldom more than a minor component of the floating weed mass when<br />
either Eichhornia crassipes or Salvinia molesta or both are present. However, when it<br />
occurs alone or when the strong competition from these two weeds is greatly reduced by<br />
their effective biological control, water lettuce can increase rapidly to occupy the vacated<br />
water surface. Since damaging biological control agents are available for all three weeds,<br />
it is sensible to embark on a biological control program for them all, either at the same<br />
time or in sequence.<br />
Adequate biological control of water lettuce has been achieved by the introduction<br />
of the weevil Neohydronomus alone. However, if an even greater degree of control is<br />
desired, there are, in South America, additional species of weevil and also several moths
4.24 Pistia stratiotes 207<br />
that appear to be well worthwhile investigating further. This is particularly so, as some<br />
are known to be heavily attacked by natural enemies and, if introduced without these,<br />
would be expected to be even more effective.<br />
It can be concluded with some confidence that water lettuce is a promising candi-<br />
date for biological control.
208 Biological Control of Weeds: Southeast Asian Prospects<br />
Portulaca oleracea<br />
(after Holm et a/. 1977)
Map 4.25 Portulaca oleracea<br />
4.25 Portulaca oleracea 209<br />
Portulaca oleracea is a serious weed throughout tropical, subtropical and temperate<br />
areas, attaining high overall pest status more because of its very widespread importance<br />
than by being amongst the top few weeds in any one country.<br />
About 14 of the 140 or so species of insects that are known to attack it appear to<br />
be restricted to the genus Portulaca and probably several to P. oleracea or its very close<br />
relatives. In their native ranges 4 leaf-mining or gall-forming flies, 1 leaf-mining moth, 1<br />
leaf-mining sawfly and 2 weevils all cause considerable damage and show high specifici-<br />
ty to P. oleracea.<br />
If this group of phytophagous insects is not already present, the establishment of<br />
several without their own natural enemies should lead to a significant lowering in the<br />
weed status of I? oleracea.<br />
Portulaca oleracea is an attractive target for an attempt at biological control.
210 Biological Control of Weeds: Southeast Asian Prospects<br />
4.25 Portulaca oleracea L.<br />
Portulacaceae<br />
pigweed, purslane; gelang, krokot (Indonesia); gelang pasir, segan (Malaysia);<br />
phak bia yai (Thailand), mya byit, mye byet (Myanmar); kbet choun (Cambodia);<br />
golasiman, ulasiman (Philippines), rau Sam (Vietnam)<br />
Rating<br />
+++ Phil<br />
10 ++ Thai<br />
+ Myan, Viet, Msia, Sing, Indo<br />
Much of the material in this dossier is summarised from the account in Waterhouse<br />
(1993b).<br />
Origin<br />
Uncertain; possibly Central America, but see comment.<br />
Distribution<br />
Very widespread in tropical, subtropical and temperate regions of the world, including<br />
Southeast Asia, Australia and the Pacific.<br />
Characteristics<br />
Pigweed is a fleshy annual herb, reproducing by seed, or by stem-fragments rooting<br />
when lying on moist soil. The stems are succulent, often reddish and 0.2 to 0.5 m in<br />
length. The stems and leaves are smooth and fleshy and form mats. In sunlight the plants<br />
are prostrate, but in partly shaded situations they may attain a height of 0.5 m. The leaves<br />
are alternate, flowers are yellow, sessile, self-pollinated and either occur singly, or sever-<br />
al may occur together, in the leaf clusters at the ends of branches. They open on sunny<br />
mornings and later produce numerous tiny (0.5 mm diameter) black seeds.<br />
Importance<br />
P. oleracea was ranked 9th of the world's worst weeds, being recorded in 45 crops in 81<br />
countries (Holm et al. 1977). With a rating of 10 in Southeast Asia, it ranked equal 32nd<br />
in the region; also 6th in the Pacific and 49th in Australia (Waterhouse 1993a,b). In<br />
Southeast Asia, it is particularly important in many upland crops, including vegetables,<br />
rice, maize, sorghum, groundnuts and sugarcane. It is drought hardy, colonising waste<br />
places and bare areas but thrives in moist fertile soils. There are many ecological types,<br />
some of which are occasionally used as a vegetable. In the Philippines up to 10000 and<br />
in Norih America up to 243 000 seeds are produced per plant. The tiny seeds are spread<br />
by wind, water, as a contaminant of the seeds of crops and by birds, surviving passage<br />
through the digestive tract. They also survive burial for long periods and germinate best<br />
above 30°C, but poorly below 24°C.
4.25 Portulaca oleracea 211<br />
Pigweed does not compete well with other weeds. It is successful because it estab-<br />
lishes rapidly after soil disturbance and may flower and seed before being outcompeted<br />
by taller plants. The succulent leaves and stems are rich in oxalates and nitrates and have<br />
been implicated in livestock deaths. The succulent leaves of some strains have been used<br />
as human food (Miyanishi and Cavers 1980).<br />
Natural enemies<br />
The 138 insect species that have been recorded attacking P. oleracea were listed by<br />
Waterhouse (1993b), most of them from Central and South America (Bennett and<br />
Cruttwell 1972) and USA (Romm 1937). A few additional records are now listed in table<br />
4.25.1. Most of these species are known to be (or suspected of being) polyphagous and<br />
many are pests. Nevertheless, table 4.25.2 lists 14 insects that, so far as known, are<br />
restricted to P. oleracea, or at least to the genus Portulaca. Eight of these appear to have<br />
originated in the Americas, 2 each in Africa and India, and 1 each in France and<br />
Southeast Asia. With the exception of the weevil Ceutorhynchus portulacae, described<br />
from P. oleracea in Java, there do not appear to be any reports of native insects which<br />
might possibly be restricted to pigweed in Southeast Asia, the Pacific or Australia.<br />
Table 4.25.1 Natural enemies of Portrclaca oleracea: additional insect records to those<br />
of Waterhouse (1 993b).<br />
Species Country Other hosts References<br />
Hemiptera<br />
APHlDlDAE<br />
Myzus persicae<br />
ClCADELLlDAE<br />
Circulifer haematoceps<br />
Orosius orientalis<br />
(= 0. albicinctus)<br />
LYGAEIDAE<br />
Nysius cymoides<br />
Nysius vinitor<br />
Coleoptera<br />
CURCULIONIDAE<br />
Hypurus bertrandi<br />
Diptera<br />
AGROMYZIDAE<br />
Liriomyza caulophaga<br />
Liriomyza trifolii<br />
1<br />
Australia polyphagous author's record<br />
Israel polyphagous Klein & Raccah 1991<br />
India polyphagous Kooner & Deol 1982<br />
Italy jojoba Parenzan 1985<br />
Australia polyphagous Elshafie 1976, Ramesh &<br />
Laughlin 1984<br />
Australia, India, specific R.E. McFadyen pers. comm.<br />
Guam, Northern 1993,Zaka-ur-rab 199 1,<br />
Marianas Zimmerman 1957<br />
Australia Beta vulgaris R.E. McFadyen pers. comm.<br />
var. cicla 1993<br />
USA polyphagous Chandler & Chandler 1988<br />
(continued on next page)
212 Biological Control of Weeds: Southeast Asian Prospects<br />
Table 4.25.1 (continued)<br />
Species Country Other hosts References<br />
Lepidoptera<br />
NOCTU IDAE<br />
Agrotis ipsilon India polyphagous Das & Ram 1988<br />
Neogalea (= Spodoptera)<br />
sunia Nicaragua polyphagous Savoie 1988<br />
Spodoprera eridania Nicaragua polyphagous Savoie 1988<br />
Spodoptera exigua Nicaragua polyphagous Savoie 1988<br />
PY RALl DAE<br />
Loxostege bijdalis N. America cotton Allyson 1976<br />
The host specificity has been investigated of 5 of the 6 species of Diptera,<br />
Lepidoptera and Hymenoptera listed in table 4.25.2, but, except for Baris arctithorax and<br />
Hypurus bertrandi, both of which appear to be adequately specific (see later), little is<br />
known about that of the 8 weevils.<br />
Several of the fungi listed in table 4.25.3 are reported to damage P. oleracea, some-<br />
times severely (Waterhouse 1993b), but too little is known about host specificity or host<br />
specific strains to evaluate their possible role as classical biological control agents.<br />
Nevertheless their specificity certainly merits investigation should the need arise.<br />
Attempts at biological control<br />
There have been no attempts to establish natural enemies as biological control agents for<br />
P. oleracea. However three insect species have appeared in countries well out of their<br />
native range, in particular the European weevil Hypurus bertrandi, but also the American<br />
sawfly Schizocerella pilicornis and the American leaf-mining fly Haplopeodes palliatus.<br />
These successful, unassisted establishments suggest that there should be little difficulty<br />
in securing assisted establishments elsewhere. Unfortunately there is no information<br />
available on what effects, if any, these three insects have had on P. oleracea in their new<br />
countries, but it is suspected that a group of species may be required to secure substantial<br />
effects in Australia.<br />
The sawfly Schizocerella pilicornis appeared in eastern Australia (Queensland and<br />
New South Wales) in the early 1960s; (Benson 1962, Krombein and Burks 1967) but<br />
there are no records of it building up in sufficient numbers to cause serious damage. In<br />
1993 Hypurus bertrandi and Liriomyza caulophaga were bred from P. oleracea leaves in<br />
Brisbane (R.E. McFadyen pers. comm.), but numbers were too low to cause serious dam-<br />
age. It is not known whether these species have only become established recently.<br />
L. caulophaga was previously known only from Beta vulgaris var. cicla (silverbeet) in<br />
Australia. Larvae tunnel in the soft white spongy tissue between the vascular strands in<br />
the leaf petioles and midribs and pupate there (Kleinschmidt 1960, 1970, Spencer 1990).<br />
EGYPT<br />
Tawfik et al. (1976) recorded Hypurus bertrandi attacking P. oleracea.<br />
GUAM AND THE NORTHERN MARIANA IS.<br />
Zimmerman (1957) records Hypurus bertrandi from Guam, Tinian, Saipan and Agrihan,<br />
some of the specimens from Saipan being taken from the crops of swifts.
4.25 Portulaca oleracea 213<br />
Table 4.25.2 Insects restricted to R oleracea or at least to the genus Portulaca (after<br />
Waterhouse 1993b).<br />
Species Distribution References<br />
Coleoptera<br />
CURCULlONlDAE<br />
Apion sp.<br />
Baris arctithorax<br />
Baris lorata<br />
Baris portulacae<br />
Ceutorhynchus oleracae<br />
Ceutorhynchus portulacae<br />
Hypurus bertrandi<br />
Linogeraeus (= Centrinaspis)<br />
perscitus<br />
Diptera<br />
AGROMYZIDAE<br />
Haplopeodes palliatus<br />
ANTHOMYIIDAE<br />
Pegornya dolosa<br />
CEClDOMYllDAE<br />
Asphondylia portulacae<br />
Lasioptera portulacae<br />
Neolasioptera portulacae<br />
Lepidoptera<br />
HELlODlNlDAE<br />
Heliodines quinqueguttata<br />
Hymenoptera<br />
TENTHREDINIDAE<br />
Schizocerella pilicornis<br />
Brazil<br />
Egypt<br />
Sudan<br />
India<br />
Java<br />
India<br />
Puerto Rico,<br />
France,<br />
Egypt,<br />
USA, Hawaii, Marianas,<br />
Australia<br />
Colombia, Trinidad, USA<br />
Australia, USA<br />
Trinidad<br />
El Salvador, Argentina,<br />
Colombia, Bolivia,<br />
Leeward Is, St Kitts<br />
Nevis, Montserrat, Jamaica<br />
USA<br />
Cuba, Florida, St Vincent<br />
Trinidad, St Kitts Nevis,<br />
Trinidad,Montserrat<br />
Puerto Rico<br />
California, Mexico<br />
USA, Australia<br />
Argentina to USA<br />
d'Araujo e Silva et al. 1968a,b<br />
Tawfik et al. 1976<br />
Marshall 19 1 1<br />
Marshall 19 16<br />
Marshall 1935<br />
Marshall 1916<br />
Wolcott 1948<br />
Tempsre 1943<br />
Tawfik et al. 1976<br />
Clement and Norris 1982<br />
RE McFadyen pers. comm. 1993,<br />
Zimmeman 1957<br />
Bennett and Cruttwell 1972,<br />
Romm 1937<br />
R.E. McFadyen pers. comm.<br />
1993, Romm 1937<br />
Bennett and Cruttwell 1972<br />
Cruttwell and Bennett 1972a<br />
Gagnt 1968,<br />
Bennett and Cruttwell 1972<br />
Felt 191 1<br />
GagnC 1968<br />
Bennett and Cruttwell 1972<br />
Bennett and Cruttwell 1972,<br />
Cruttwell and Bennett 1972b<br />
Wolcott 1948<br />
Bennett and Cruttwell 1972<br />
Krombein and Burks 1967<br />
Muesebeck et al. 195 1
214 Biological Control of Weeds: Southeast Asian Prospects<br />
Table 4.25.3 Natural enemies of Portulaca oleracea: fungi, viruses and nematodes.<br />
Species Country Other hosts References<br />
FUNGI<br />
AIbugo portulacae<br />
AIbugo portulacearum<br />
Ascochyta portulacae<br />
Bipolaris (= Drechslera)<br />
indica<br />
Cercospora portulacae<br />
Cercosporella dominicana<br />
Dendrographium<br />
lucknowense<br />
Dichotomophthora indica<br />
(= D. lutea)<br />
Dichotomophthora<br />
portulacae<br />
Helminthosporium<br />
(Bipolaris) portulacae<br />
Phoma sp.<br />
Phytophthora palmivora<br />
Polymyxa betae f. sp.<br />
portulacae<br />
VIRUSES<br />
anemone brown ring<br />
aster yellows<br />
beet curly top<br />
chili vein banding<br />
clover big vein<br />
cucumber mosaic<br />
groundnut rosette<br />
tobacco broad ring spot<br />
tobacco etch<br />
tobacco mosaic<br />
tobacco streak<br />
NEMATODES<br />
Criconemella xenoplax<br />
Ditylenchus dipsaci<br />
Helicotylenchus indicus<br />
Helicotylenchus<br />
multicinctus<br />
Heterodera glycines<br />
Europe, Africa,<br />
Asia, Americas<br />
Poland<br />
USSR<br />
USA<br />
India<br />
Dominica<br />
India<br />
IMI 1992, Miyanishi &<br />
Cavers 1980<br />
IMI 1992<br />
IMI 1992<br />
many, including Evans 1987,<br />
Amaranthus spinosus Kenfield et al. 1989<br />
IMI 1992<br />
IMI 1992<br />
IMI 1992<br />
USA, West Indies, Helianthus,<br />
Europe, India Pennisetum<br />
Europe, Sudan, tarragon, cactus,<br />
Hawaii, capsicum,<br />
California, Glycine max<br />
Jamaica<br />
Venezuela<br />
USA Portulaca grandifora<br />
France tarragon<br />
Sarawak pepper and several<br />
weeds<br />
Bulgaria, Japan sugar beet, wheat,<br />
many weeds<br />
Hawaii<br />
Hawaii<br />
Hawaii<br />
Hawaii<br />
Hawaii<br />
Bulgaria<br />
Hungary<br />
USA<br />
Malawi<br />
Hawaii<br />
Hawaii<br />
Philippines<br />
Hawaii<br />
anemone<br />
aster<br />
sugar beet<br />
chili<br />
clover<br />
many economic<br />
plants<br />
cucumber, tobacco<br />
groundnut<br />
tobacco<br />
tobacco<br />
many<br />
tobacco<br />
Baudoin 1986, IMI 1992,<br />
Rao 1966<br />
Vegh & Le Berre 1984<br />
Klisiewicz 1985,<br />
Klisiewicz et al. 1983,<br />
Mehrlich & Fitzpatrick 1935<br />
Mitchell 1986<br />
IMI 1992, Rader 1948,<br />
Strider & Chi 1984<br />
Vegh & Le Berre 1984<br />
Anon 1979<br />
Abe & Ui 1986,<br />
Vrbanov & Krumov 1989<br />
Holm et al. 1977<br />
Holm et al. 1977<br />
Holm et al. 1977<br />
Holm et al. 1977<br />
Holm et al. 1977<br />
Dikova 1989,<br />
Nasser & Basky 1988,<br />
Dodds & Taylor 1980<br />
Adams 1967<br />
Holm et al. 1977<br />
Holm et al. 1977<br />
Eugenio & del Rosario 1962<br />
Holm et al. 1977<br />
USA many legumes and Zehr et al. 1990<br />
other plants<br />
USSR polyphagous Kholod 1983<br />
India polyphagous Rahman & Khan 1986<br />
Brazil, Ivory banana Luc et al. 1990,<br />
Coast Zem & Lordello 1983<br />
Colombia soybean Quintero et al. 1988<br />
I (continued on next page)
4.25 Portulaca oleracea 215<br />
Species Country Other hosts References<br />
Heterodera rnarioni<br />
Hoplolairnus indicus<br />
Meloidogyne sp.<br />
Meloidogyne arenaria<br />
Meloidogyne hapla<br />
Meloidogyne incognita<br />
Meloidogyne javanica<br />
Pratylenchus rninutus<br />
Pratylenchus sp.<br />
Radopholus sirnilis<br />
Rotylenchulus reniforrnis<br />
Tylenchorhynchus<br />
brassicae<br />
Hawaii<br />
USA<br />
Cuba<br />
USA<br />
Hungary<br />
India, USA,<br />
Philippines<br />
India<br />
Hawaii<br />
Ivory Coast<br />
Ivory Coast<br />
India, USA,<br />
Hawaii<br />
India<br />
eggplant, tomato<br />
coffee<br />
tobacco<br />
several weeds<br />
polyphagous<br />
polyphagous<br />
Musa sp.,<br />
several weeds<br />
ornamentals and<br />
many weeds<br />
polyphagous<br />
Linford & Yap 1940<br />
Rahman & Khan 1986<br />
Izquierdo et al. 1987<br />
Tedford & Fortnum 1988<br />
Dabaj & Jenser 1990<br />
Maqbool et al. 1986,<br />
Tedford & Fortnum 1988,<br />
Valdez 1968<br />
Maqbool et al. 1986<br />
Linford et al. 1949<br />
Luc et al. 1990<br />
Luc et al. 1990<br />
Heald et al. 1974,<br />
Inserra et al. 1989<br />
Khan & Khan 1985<br />
Linford & Yap 1940<br />
Rahman & Khan 1986<br />
HAWAII<br />
Pigweed was established in Hawaii prior to 187 1 (Hillebrand 1888). Hypurus bertrandi,<br />
originally misidentified as Ceutorhynchus sp., was reported in 1950 to be numerous<br />
enough in many cases to defoliate P. oleracea and to cause it to collapse as if sprayed<br />
with a herbicide (Bianchi 1955). Nevertheless, in 1992, Hawaiian weed scientists considered<br />
it as one of their worst weeds (W.C. Mitchell pers. comm. 1992), so the control<br />
exerted by H. bertrandi and v'arious non-specific insects is clearly insufficient.<br />
Zimmerman (1957) postulates that H. bertrandi was introduced from the Marianas to<br />
Hawaii, possibly with war material being returned from the battlefields. However it is<br />
more likely to have moved in the reverse direction.<br />
INDIA<br />
Zaka-ur-rab (1991) records Hypurus bertrandi as one of the leaf-mining weevils of the<br />
Indian subcontinent.<br />
PUERTO RlCO<br />
Wolcott (1948) recorded H. bertrandi from Puerto Rico, but no other information is<br />
available.<br />
Major Natural Enemies<br />
Summarised below is what is known of the biology of nine of the natural enemies listed<br />
in table 4.25.2.<br />
Apion sp. Coleoptera: Curculionidae<br />
Apion sp. produces galls in the flower buds of P. oleracea in Brazil (d'Araujo e Silva et<br />
al. 1968a,b) and Apion larvae also cause significant damage by gall formation in flowers<br />
in northern Argentina (Bennett and Cruttwell 1972, Bennett pers. comm. 1992).
216 Biological Control of Weeds: Southeast Asian Prospects<br />
Asphondylia portulacae Diptera: Cecidomyiidae<br />
Oviposition by this flower gall midge into the very small pigweed buds causes them to<br />
develop abnormally. Usually only one larva develops per bud and occupies a chamber in<br />
the swollen receptacle. Galled flowers do not produce seed. A. portulacae is heavily<br />
attacked by parasitoids (Bennett and Cruttwell 1972) and might be an important natural<br />
enemy if freed from them. The genus Asphondylia is considered to be highly host specif-<br />
ic and 52 of its 54 species are known from only a single host. Each of the two exceptions<br />
only attacks two plants of the same genus and Bennett and Cruttwell (1972) suggested,<br />
on the basis of this information, that host specificity testing was unnecessary.<br />
Baris arctithorax Coleoptera: Curculionidae<br />
In Egypt this weevil forms stem galls on pigweed, but is not known from any economic<br />
plant. Eggs are laid singly in cavities gnawed in the stem by the female, leading to the<br />
production of single closed galls in which the larvae feed. Pupation occurs in the soil.<br />
Young infested plants produce weak vegetative growth, few seeds and may even be<br />
killed. Adult weevils feed on the surface of leaves causing white blotches or holes. At<br />
temperatures from 25 to 30°C the development time from egg to adult is about 40 days.<br />
Infestations of up to 74% of plants are recorded in summer and a peak of 95% in autumn<br />
(Awadallah et al. 1976, Tawfik et al. 1976).<br />
Haplopeodes palliatus Diptera: Agromyzidae<br />
The genus Haplopeodes contains 13 species, all from the Americas and known on only<br />
four plant families-Portulacaceae (1 species) Amaranthaceae (3 species),<br />
Chenopodiaceae (2 species) and Solanaceae (8 species) (Spencer 1990). Each appears to<br />
be specific to a single genus and H. palliatus is known only from l? oleracea. It is a typi-<br />
cal leaf miner.<br />
Heliodines quinqueguttata Lepidoptera: Heliodinidae<br />
Eggs, which are laid singly or in groups of up to 6, hatch in 5 to 6 days and larvae wan-<br />
der some distance over the plant before mining into a leaf, stem or seed capsule. After 7<br />
to 8 days the fifth instar larva leaves the mine and pupates within a flimsy silk cocoon<br />
attached to the stems or leaves of the plant. Larvae are attacked by a braconid endopara-<br />
sitoid, Pholetesor (= Apanteles) sp. (circumscriptus group).<br />
Host specificity tests were carried out on a wide variety of economic and non-<br />
economic plants, but development was completed only on Portulaca oleracea, P. pilosa<br />
(also weedy) and the ornamental P. grandiflora. However, in the field in Trinidad, neither<br />
P. pilosa nor P. quadriJida (also weedy) were attacked and l? grandiflora was not grown.<br />
There appear to be no records of Heliodines species attacking crops and each species<br />
appears to be restricted to a single plant family. Cruttwell and Bennett (1972b) suggested,<br />
therefore, that it should be considered as a biological control agent.<br />
Hypurus bertrandi Coleoptera: Curculionidae<br />
This tiny (2mm long) weevil has spread unaided from its native France to Egypt (prior to<br />
1926) (Hoffman and Tempbre 1944, Tawfik et al. 1976), Puerto Rico (Wolcott 1948),<br />
Hawaii (1950) (Davis 1955, Maehler 1954), Guam and the Northern Marianas
4.25 Portulaca oleracea 217<br />
(Zimmerman 1957), California (1980) and Queensland (1993) (R.E. McFadyen pers.<br />
comm.).<br />
Eggs are deposited singly and larvae mine the leaves. Infested leaves wilt and fall;<br />
and the larvae then migrate to fresh leaves, each destroying four or five in its lifetime. If<br />
no undamaged leaves are available the outer tissues of stems are attacked. Pupation<br />
occurs in a cell formed by soil particles cemented by fecal secretion and, in France,<br />
adults overwinter under the bark of trees. Adults feed on leaf margins, stems and devel-<br />
oping seed capsules. Development is rapid, from egg to adult in 10 days at 32.2OC and<br />
under 16 hours light. Z? oleracea is its only reported host plant. In France it is parasitised<br />
by a number of wasps (Tawfik et al. 1976, Clement and Norris 1982, Hoffmann and<br />
Tempbre 1944, Norris 1 985, Tempbre 1943, 1 944, 1950).<br />
Neolasioptera portulacae Diptera: Cecidomyiidae<br />
Oviposition in the stem by this midge leads to globular galls up to 1.5 cm in diameter,<br />
each containing up to 10 larvae. Galls retard or prevent growth and also flower and seed<br />
production. In open, infertile sites every pigweed stem may be infested but, in vigorous<br />
growth or in shaded sites, the level of attack is usually very low. The larvae are heavily<br />
attacked by parasitoids.<br />
All except one of the 51 species of Neolasioptera are restricted to one plant genus<br />
and the remaining species only attacks two plant genera. This was taken by Bennett and<br />
Cruttwell (1972) to indicate that N. portulacae is sufficiently host specific to be<br />
employed for biological control.<br />
Pegomya dolosa Diptera: Anthomyiidae<br />
Eggs are laid singly on the underside of the pigweed leaf and hatch after about 3 days.<br />
The larvae are leaf miners, devouring the contents of the leaf and then leaving to enter<br />
another. Two or more leaves are commonly destroyed by each larva. After feeding for<br />
about 7 days larvae leave to pupate in the soil, later emerging as 3 to 4 mm long adults.<br />
Eggs are parasitised and larvae are attacked by a pteromalid wasp.<br />
Schizocerella pilicornis Hymenoptera: Tenthredinidae<br />
This leaf-mining sawfly occurs naturally over a very wide range from Argentina and<br />
Brazil to USA (Muesebeck et al. 195 1). It appeared unaided in eastern Australia (Benson<br />
1962, Krombein and Burks 1967). There are two biotypes. The larvae of one which is<br />
widespread in USA mines the leaves, whereas those of the other (from Mississippi north-<br />
wards) feeds externally on the leaves.<br />
Females mate soon after emergence and lay up to 40 eggs singly in the edges of the<br />
leaves. The larvae mine the leaves, damaging each to the point of collapse before moving<br />
to another. At least two leaves are destroyed by each larva. The mature larvae enter the<br />
soil and spin cocoons. The life cycle can be completed in 13 days and there are a number<br />
of generations each year (Clement and Norris 1982, Gorske et al. 1976). In California<br />
prepupae in diapause overwinter in the soil. Adults live for a day and do not feed. In<br />
California up to 84% of Z? oleracea leaves were severely damaged, leading to defoliation<br />
and sometimes death of the plant. When Z? oleracea was protected by insecticide from<br />
both S. pilicornis and Hypurus bertrandi it produced about four times as much seed as
218<br />
Biological Control of Weeds: Southeast Asian Prospects<br />
unprotected plants, although the latter still produced enough (4000 to 5000/m2/day) to<br />
maintain a high seed bank in the soil (Force 1965, Garlick 1922, Gomes de Lima 1968,<br />
Gorske et al. 1977, Norris 1985, Webster and Mally 1900).<br />
S. pilicornis has not been recorded from any economic plant and, in starvation tests,<br />
was only able to feed on P. oleracea and the related Montia perfoliata (Gorske et al.<br />
1976). A microsporidan, Nosema pilicornis causes high mortality in infected S. pilicornis<br />
larvae in USA and should be eliminated during any transfer of the sawfly to new areas<br />
(Gorske and Maddox 1978).<br />
Comment<br />
It is generally believed that, through coevolution, there are likely to be a number of spe-<br />
cific (or nearly specific) natural enemies of an organism in its area of origin.<br />
Furthermore, that not all such organisms will have accompanied their host when it has<br />
moved outside its area of origin. If this generalisation is applied to P. oleracea it can be<br />
seen that 8 of its 14 specific (or nearly specific) natural enemies are of American origin<br />
but no more than 2 from any other region of the world (Table 4.25.2). The inference from<br />
this is that P. oleracea is probably of American origin, an inference supported by the<br />
finding of seeds in Louisiana, Illinois and Kentucky dating between lOOOBC and 750AD<br />
and pollen and seeds in Ontario sediments dating back to 1350AD (Miyanishi and Cavers<br />
1980). This suggests that, if it is desired to evaluate insects additional to those listed in<br />
table 4.25.2, they should first be sought from the Americas and possibly from amongst<br />
those listed by Waterhouse (1993b).<br />
The family Portulacaceae is relatively small with 20 genera and about 250 species<br />
worldwide, of which the genus Portulaca contains about half (West 1990). Very few<br />
Portulacaceae are cultivated: Portulaca grandiflora as a brightly flowering garden plant,<br />
Talinum paniculatum and T. triangulare as pot herbs (but they are sometimes regarded as<br />
weeds), Montia fontana for salads, Lewisia spp. as rockery plants and Anacampseros as a<br />
succulent; but these are not considered to be of great economic importance (Cruttwell<br />
and Bennett 1972a).<br />
From what is known about the biology of the insects in table 4.25.2 it appears that,<br />
if a group of them is established in a new country without their own natural enemies,<br />
they should cause serious damage to l? oleracea and significantly reduce its competitive-<br />
ness and seed production.
220 Biological Control of Weeds: Southeast Asian Prospects<br />
Rottboellia cochinchinensis<br />
(after Holm eta/. 1977)
Map 4.26 Rottboellia cochinchinensis<br />
4.26 Rottboellia cochinchinensis 221<br />
Rottboellia cochinchinensis<br />
Rottboellia cochinchinensis is probably of Southeast Asian or Indian origin, although one<br />
biotype may have evolved in East Africa. Very little is known of its insect enemies in<br />
these regions and they certainly merit investigation. Current research on fungal<br />
pathogens indicates that Sporisorium ophiuri is a potential agent for the Americas and<br />
other regions where it does not yet occur.<br />
If preliminary surveys for specific insect enemies in Asia and Africa support a<br />
Southeast Asian origin it is unlikely to be an attractive early target for classical biological<br />
control in this region.
222 Biological Control of Weeds: Southeast Asian Prospects<br />
4.26 Rottboellia cochinchinensis (Lour.) W.D. Clayton<br />
(= Rottboellia exaltata)<br />
Poaceae<br />
itch grass; myet ya, myet ya nge (Myanmar), branjangan (Indonesia), aguitigay<br />
(Philippines), yaa prong khaai (Thailand)<br />
Rating<br />
+++ Indo, Phil<br />
12 ++ Thai<br />
+ Myan, Laos, Viet, Msia<br />
Origin<br />
Uncertain but probably India to Southeast Asia; one biotype in East Africa.<br />
Distribution<br />
Worldwide, and weedy between 23' north and south latitudes (Holm et al. 1977).<br />
Formerly known as Rottboellia exaltata. Two biotypes have been recognised in USA and<br />
at least five in the Philippines (Fisher et al. 1987). The origin of R. cochinchinensis has<br />
not been completely resolved. It is said to be native to India (Holm et al. 1977), but in his<br />
survey of fungal pathogens and their specificity (or lack of it) Evans (pers. comm. 1992)<br />
was led to the conclusion that it may be Southeast Asia; and furthermore, that there is a<br />
distinct East African biotype, with possibly specific natural enemies of its own. It is<br />
interesting that, of the 14 Kenyan tribes visited by Ellison and Evans (1990), 13 had a<br />
separate and specific vernacular name for itch grass. Although it was a common weed<br />
and dense stands occurred at the edge of fields and along roadsides, control was achieved<br />
by early season hoeing and none of the farmers suggested it was a major problem. This<br />
evidence was taken as reinforcing the theory that East Africa is the centre of origin at<br />
least of the local biotype of the weed.<br />
Characteristics<br />
R. cochinchinensis is a tall, erect, strongly tufted, annual grass growing to 3 m. It has stilt<br />
roots. Its leaves and stems have long, sharp, fragile, siliceous, irritating hairs that break<br />
off in the flesh on contact. The inflorescence is a single cylindrical spike.<br />
Importance<br />
It is an aggressive C4, annual grass of 18 tropical and subtropical crops in 28 countries,<br />
including maize, rice, sorghum, soybeans and sugarcane. The heaviest infestations occur<br />
in the Caribbean, Central America, and parts of South America, to which it is a relatively<br />
recent introduction; also a widespread weed in southern Africa. It is an important weed in<br />
sugarcane, maize and upland rice in the Philippines. It is often a primary coloniser of dis-<br />
turbed land. It flowers all year round. Reproduction is by seed and up to 8000 may be
4.26 Rottboellia cochinchinensis 223<br />
produced per plant. The seed is about the size of a rice grain and is not easily separated<br />
from intermingled rice grains. Some seeds germinate immediately, whereas others lie<br />
dormant for varying periods, sometimes years. Buried seeds may germinate and emer-<br />
gence take place from a depth of 15 cm. It is common in open, well-drained places, but<br />
also grows in wet places and even in shallow water. It commonly occurs on contour<br />
banks and roadsides. It prefers sunny or no more than moderately shaded situations, but<br />
can grow in deep shade. Its many needle-like hairs deter hand removal of older plants,<br />
since the hairs penetrate hands and clothing and result in painful infections.<br />
R. cochinchinensis is sometimes grazed and used for green fodder, although avoided<br />
at times by some animals because of its sharp hairs.<br />
Natural enemies<br />
Fungi are the only natural enemies (Tables 4.26.1 to 4.26.3) for which there is any rea-<br />
sonably comprehensive information. Fungi have been surveyed and their specificity is<br />
being examined in a joint International Institute of Biological Control and Long Ashton<br />
Research Station project covering East Africa, South America, India, Nepal, Sri Lanka<br />
and Thailand (Ellison 1992, Ellison and Evans 1990, 1993, Evans 1991, Natural<br />
Resources Institute 1992).<br />
As indicated earlier, R. cochinchinensis shows high biotype variation between coun-<br />
tries and this is correlated with varying levels of susceptibility to different fungal isolates.<br />
Also, a clear positive correlation was found between high pathogen virulence and inade-<br />
quate specificity to itch grass (Ellison 1992). Maize (Zea mays) proved to be the crop<br />
species most at risk from itch grass pathogens, which is not surprising in view of the<br />
close evolutionary relationship of the two genera involved. This suggests that the use of<br />
fungi for classical biological control of itch grass may not show great promise, but that<br />
their use as mycoherbicides might prove effective. All except one of the fungi from tropi-<br />
cal America that have been tested are non-specific to itch grass and most are local<br />
pathogens that have transferred from local grasses (Evans 1987). In Kenya, in addition to<br />
at least 10 non-specific fungi attacking itch grass, a head smut, Sporisorium<br />
(= Sphacelotheca) ophiuri was found, which appears to be restricted to Rottboellia and<br />
the closely related genus Chasmopodium (Ellison and Evans 1990, Zundel 1953).<br />
S. ophiuri is recorded as occurring in East Africa, Sri Lanka, Philippines and<br />
Thailand, but apparently not in the Americas. It is often locally damaging, significantly<br />
reducing vigour and virtually eliminating seeding. Its host specificity is under detailed<br />
investigation (Ellison and Evans 1993, Evans 1991) as a potential candidate for classical<br />
biological control for areas where it does not already occur. In an annual weed where<br />
seeds are the only means of propagation, a destructive seed head disease, such as S. ophi-<br />
uri, is a highly promising biological control agent (Evans 1991).<br />
A Curvularia isolate from Trinidad proved highly damaging to itch grass, while not<br />
damaging rice, maize, sugarcane or pearl millet (Evans 1991). A Curvularia from<br />
Somalia was able to kill R. cochinchinensis in a few days, but was also able to infest<br />
maize. However the crop readily recovered (Ellison 1992). If further tests confirm its<br />
specificity, it may be a potential biological control agent. The same applies to Puccinia
224 Biological Control of Weeds: Southeast Asian Prospects<br />
rottboelliae about which less is recorded (Evans 1987). Special attention is now being<br />
paid to the possibility of developing preparations of one or more of these fungi as a<br />
mycoherbicide. An isolate from Thailand of Colletotrichium sp. which appears to be spe-<br />
cific to itch grass has been selected from 900 fungal samples and field trials have already<br />
demonstrated that an appropriate formulation has potential against this weed, particularly<br />
when combined with low doses of herbicide (Ellison 1992, Ellison and Evans 1993,<br />
Natural Resources Institute 1992).<br />
Surprisingly few insects (Table 4.26.1) have been recorded attacking itch grass and<br />
only one unidentified gall midge recorded in India from Rottboellia compressa (Barnes<br />
1946) might, if it attacks R. cochinchinensis also, be specific enough to be a candidate<br />
agent. It is regrettable that parallel observations were not made on insects during the<br />
extensive fungal surveys. In East Africa a stem borer, a lepidopteran leaf feeder and a fly<br />
larva all proved to be non-specific graminaceous feeders (H.C. Evans pers. comm. 1992,<br />
1993).<br />
R. cochinchinensis is an alternative host for a number of viruses, almost all of them<br />
serious diseases of maize (Table 4.26.3). It is surprising that the only record encountered<br />
dealing with nematodes related to a study of 16 plant parasitic species attacking sugar-<br />
cane in the Philippines. This found that itch grass was not infected by Meloidogyne sp.<br />
(Reyes and Beguico 1978).<br />
Comment<br />
Rottboellia belongs to the same grass tribe (Andropogoneae), but not to the same sub-<br />
tribes, as Saccharum, Sorghum and Zea (Table 4.26.4). This suggests that candidates for<br />
classical biological control of this weed will have to pass extensive host specificity test-<br />
ing against all of the crop and pasture grasses belonging to these and related genera<br />
before being considered for release.<br />
Table 4.26.1 Natural enemies of Rottboelliu cochinchinensis: insects.<br />
Species Location Other hosts References<br />
Orthoptera<br />
GRYLLIDAE<br />
Euscyrtus concinnus Philippines Eleusine indica, Barrion & Litsinger 1980<br />
Dactylocteniurn<br />
aegyptiurn, Cyperus<br />
rotundus, Digitaria<br />
sanguinalis<br />
Pteronernobius sp. Sierra Leone Alghali & Domingo 1982<br />
Hemiptera<br />
APHlDlDAE<br />
Myzus persicae Peru a very wide range Ortiz 198 1<br />
Uroleucon (= Dactynotus)<br />
. ambrosiae USA sorghum, maize Koike 1977<br />
I<br />
(continued on next page)
4.26 Rottboellia cochinchinensis 225<br />
Species Location Other hosts References<br />
CICADELLIDAE<br />
Nephotettix nigrornaculatus Sierra Leone<br />
DELPHACIDAE<br />
Peregrinus rnaidis Venezuela<br />
Coleoptera<br />
COCCINELLIDAE<br />
Chnootriba (= Epilachna)<br />
similis Sierra Leone<br />
Diptera<br />
AGROMYZIDAE<br />
Pseudonapornyza<br />
philippinensis Philippines<br />
CEClDOMYllDAE<br />
a gall midge India<br />
MUSCIDAE<br />
Atherigona soccata Kenya<br />
Lepidoptera<br />
LYMANTRIIDAE<br />
Psalis pennatula Kenya<br />
NOCTUIDAE<br />
Sesarnia sp. Ghana<br />
Spodoptera frugiperda USA<br />
PYRALIDAE<br />
Chilo sp.<br />
sp.?<br />
Ghana<br />
Kenya<br />
rice, Ischaernurn Alghali & Domingo 1982<br />
rugosurn, Paspalurn<br />
vaginaturn<br />
a virus transmitter Ferreira et al. 1989, Migliori<br />
on many hosts & Lastra 1980, 198 1,<br />
Trujillo et al. 1974<br />
Alghali & Domingo 1982<br />
known only from Spencer 1961<br />
R. cochinchinensis<br />
recorded only from Barnes 1946<br />
Rottboellia cornpressa<br />
rice Ogwaro 1978<br />
generalist Poaceae H.C. Evans pers. comm. 1993<br />
leaf eater<br />
Sampson & Kumar 1986<br />
many graminaceous Rajapakse et al. 1988<br />
crops<br />
Sampson & Kumar 1986<br />
generalist Poaceae H.C. Evans pers. comm. 1993<br />
leaf eater<br />
Table 4.26.2 Natural enemies of Rottboellia cochinchinensis: fungi.<br />
Fungi Location Other hosts References<br />
Ascochyta sp.<br />
Bipolai-is perotidis<br />
Cercospora spp.<br />
Kenya no tests on other Ellison & Evans 1990, 1993<br />
plants<br />
Australia many other grasses QDPI, unpublished<br />
Kenya, Ethiopia, maize mildly Ellison & Evans 1990, 1993<br />
Zanzibar,<br />
Madagascar,<br />
Americas, SE Asia<br />
(continued on next page)
226 Biological Control of Weeds: Southeast Asian Prospects<br />
Table 4.26.2 (continued)<br />
Fungi<br />
Cercospora fusimaculans<br />
Cercospora rottboelliae<br />
Cochliobolus<br />
(Helminthosporiurn) bicolor<br />
Cochliobolus (Curvularia)<br />
cymbopogonis<br />
Cochliobolus heterostrophus<br />
(Drechslera maydis)<br />
Colletotrichium sp.<br />
Coniothyrium sp.<br />
Curvularia spp. (many)<br />
Diaporthe (Phomopsis) sp.<br />
Diplodia sp.<br />
Fusarium moniliforme<br />
(Gibberella fujikuroi)<br />
Glornerella (Colletotrichium)<br />
grarninicola<br />
Leptosphaeria sp.<br />
Magnaporthe (Pyricularia)<br />
grisea<br />
Phaeoseptoria sp.<br />
Phyllachora sacchari<br />
Puccinia rottboelliae<br />
Location Other hosts References<br />
Sudan, Zambia, many grasses<br />
Ghana, Guinea,<br />
Togo,.Uganda,<br />
Jamaica<br />
Guinea<br />
Zimbabwe, sugarcane, maize,<br />
Somalia pearl millet<br />
Bolivia, Kenya, sugarcane, maize,<br />
Trinidad, pearl millet,<br />
Zanzibar, USA, sorghum<br />
SE Asia<br />
Kenya, Papua sugarcane, maize<br />
New Guinea,<br />
SE Asia<br />
Thailand (this species is close<br />
to C. graminicola)<br />
Africa, SE Asia<br />
Trinidad, Papua most (not all) attack<br />
New Guinea, economic crops<br />
Madagascar,<br />
Somalia, Zanzibar<br />
Kenya, SE Asia maize mildly<br />
Kenya no tests on other<br />
plants<br />
Guatemala the particular strain<br />
tested had limited<br />
host range with no<br />
symptoms in maize,<br />
sorghum or sugar<br />
Jimenez et al. 1990<br />
cane, but other strains<br />
attack these and rice<br />
India, Nepal, No Colletotrichiurn Evans 1987, 1991<br />
Sri Lanka, infection observed<br />
Thailand in East Africa<br />
Kenya, SE Asia no tests on other<br />
plants<br />
Ellison & Evans 1990, 1993<br />
Kenya, maize mildly, also<br />
Zimbabwe Eleusine spp.<br />
Kenya, SE Asia no tests on other<br />
plants<br />
Asia, Nigeria,<br />
Sicily, Argentina<br />
Kenya, Mada- limited host range<br />
gascar, Ghana,<br />
Uganda, Zambia,<br />
Ethiopia, Guinea,<br />
Nigeria, Sierra<br />
Leone, Sudan,<br />
Zimbabwe, India<br />
Evans 1987<br />
Evans 1987<br />
Ellison & Evans 1990. 1993<br />
Ellison & Evans 1990, 1993,<br />
Evans 1987, Walker &<br />
White 1979<br />
Ellison & Evans 1990, 1993,<br />
Evans 1987<br />
Ellison 1992, Ellison & Evans<br />
1993<br />
Ellison & Evans 1993<br />
Ellison 1992, Evans 1991<br />
Ellison & Evans 1990, 1993<br />
Ellison & Evans 1990. 1993<br />
Ellison & Evans 1990, 1993,<br />
Evans 1987<br />
Ellison & Evans 1990, 1993<br />
Anahosur and Sivanesan 1978<br />
Ellison & Evans 1990, 1993,<br />
Evans 1987<br />
(continued on next page)
4.26 Rottboellia cochinchinensis 227<br />
Fungi Location Other hosts References<br />
Pyrenochaeta sp. SE Asia<br />
Sphacelotheca rottboelliae Malawi, India<br />
Sporisorium (= Sphacelotheca) Kenya, Somalia,<br />
ophiuri Sudan, Uganda,<br />
Zimbabwe,<br />
Sierra Leone,<br />
Sri Lanka,<br />
Philippines<br />
Ustilago scitaminea Philippines<br />
Ellison & Evans 1993<br />
also on Saccharum Evans 1987<br />
spontaneum<br />
limited host range, Ellison & Evans 1990, 1993,<br />
and not present Evans 1987, 1991<br />
in the Americas<br />
Latiza 1980<br />
Table 4.26.3 Natural enemies of Rottboellia cochinchinensis: viruses.<br />
Virus Location Other hosts References<br />
corn leaf gall virus Philippines Agati & Calica 1950<br />
maize stripe tenuivirus USA sorghum Bradfute & Tsai 1990<br />
maize stripe virus USA sorghum Gingery et al. 1981<br />
maize hoja blanca Venezuela sorghum Ferreira et al. 1989<br />
maize white leaf Venezuela Trujillo et al. 1974<br />
maize rayado fine Texas several grasses Nault et al. 1980<br />
maize mosaic Guadeloupe,<br />
French Guinea sorghum Migliori & Lastra 198 1<br />
maize dwarf mosaic USA sorghum, sugarcane Gillespie & Koike I973<br />
maize yellow mottle Nigeria Thottapilly et al. 1992<br />
virus like disease of maize Guadaloupe sorghum Migliori & Lastra 1980<br />
rice leaf gall Philippines Agati & Calica 1950<br />
Table 4.26.4 Taxonomic position of the major exotic grass weeds (bold text) in rela-<br />
tion to the major genera of crops in the family Poaceae (= Gramineae).<br />
Subfamily Tribe Sub-tribe Genera<br />
Bambusoideae Bambuseae<br />
Oryzeae<br />
Pooideae Triticeae<br />
Aveneae<br />
Chloridoideae Eragrostideae<br />
Cynodonteae<br />
Panicoideae Paniceae<br />
Andropogoneae<br />
Maydeae<br />
Bambusa<br />
Oryza<br />
Hordeum, Secale,<br />
Triticum<br />
Avena<br />
Eleusine, Eragrostis<br />
Cynodon<br />
Setariinae Echinochloa, Panicum,<br />
Paspalum, Setaria<br />
Digitariinae Digitaria<br />
Cenchrinae Pennisetum<br />
Saccharinae Saccharum<br />
Sorghinae Sorghum<br />
Rottboelliinae Rottboellia<br />
Tripsacinae Zea
228 Biological Control of Weeds: Southeast Asian Prospects<br />
Sphenoclea zeylanica<br />
(after Holm et a/. 1977)
Map 4.27 Sphenoclea zeylanica<br />
4.27 Sphenoclea zeylanica 229<br />
Sphenoclea zeylanica<br />
Goose weed, Sphenoclea zeyhnica, is native to Tropical Africa where it is not regarded<br />
as a weed, although there are no natural enemies recorded from it there. In Southeast<br />
Asia, where it is an important weed of rice, and in India, it is sometimes severely affected<br />
by a fungus which may have some promise as a bioherbicide.<br />
A survey for natural enemies in its area of origin would be required to evaluate<br />
the prospects for its biological control.
230 Biological Control of Weeds: Southeast Asian Prospects<br />
4.27 Sphenoclea zeylanica Gaertn.<br />
Sphenocleaceae<br />
gooseweed; goenda (Indonesia), silisilihan (Philippines), pakpawd, phak pot<br />
(Thailand) xa b6ng (Vietnam)<br />
Rating<br />
+++ Msia<br />
14 ++ Thai, Camb, Phil<br />
+ Myan, Laos, Viet, Brun, Indo<br />
Origin<br />
Tropical Africa<br />
Distribution<br />
In tropical and subtropical regions across the world. From Iran extending eastwards to<br />
Indonesia and the Philippines, also China, Japan, USA, the ~aribbean, Guyana, Surinam<br />
and Madagascar. Not reported from Papua New Guinea, Australia or the oceanic Pacific.<br />
Characteristics<br />
S. zeylanica is an erect, fleshy, herbaceous annual, often with much branched, hollow<br />
stems, growing to 1.5 m; leaves alternate, oblong to lanceolate, tapering to both ends;<br />
flowers sessile in dense spikes, terminal, whitish; seed yellowish brown, 0.5 mm long;<br />
roots cord-like.<br />
Importance<br />
S. zeylanica is unusual in that it is not reported as a weed in any crop except rice (Holm<br />
et al. 1977). It thrives in damp ground at altitudes up to 350 m. In Africa it grows in the<br />
mud of tidal creeks, but does not have this habit in Malaysia. It occurs on the sides of<br />
ponds and along ditches and rivers, on dry river beds and in seasonal swamps. It prefers<br />
stagnant water sites. It reproduces continuously by seed in the Philippines. In spite of its<br />
competition with the rice plant, S. zeylanica can give valuable practical control (up to<br />
99%) of populations of rice nematodes (Hirschmaniella spp.), with the additional benefit<br />
of increased soil nitrogen. It acts through the production of toxic plant exudates<br />
(Mohandes et al. 1981).<br />
In Java, young plants and tips of older plants are steamed and eaten with rice.<br />
Natural enemies<br />
The only natural enemy encountered in the literature search is a fungus (Table 4.27.1).<br />
This was a severe infestation of the fungus Cercosporidiurn helleri on the lower surface<br />
of S. zeylanica leaves in India (Ponnappa 1967). The affected leaves became deformed<br />
and fell off. Similar fungi capable of causing death of the weed were observed at Los
4.27 Sphenoclea zeylanica 231<br />
Banos, Philippines and at Prey Phadu, Cambodia (Moody et al. 1987). If this fungus<br />
proves to be adequately specific it may have some value as a bioherbicide.<br />
Table 4.27.1 Natural enemies of Sphenoclea zeylanica<br />
Species Location References<br />
NEMATODE<br />
Meloidogyne graminicola Luc et al. 1990<br />
FUNGUS<br />
Cercosporidium helleri Cambodia, India, Moody et al. 1987,<br />
Philippines Ponnappa 1967
5 References<br />
Abe, H. and Ui, T. 1986. Host range of<br />
Polymyxa betae Keskin strains in rhizoma-<br />
nia-infested soils of sugar beet fields in<br />
Japan. Annals of the Phytopathological<br />
Society of Japan 52: 394-403. (Weed<br />
Abstracts 36: 2228, 1987).<br />
Abenes, M.L.P. and Khan, Z.R. 1990. Feeding<br />
and food assimilation by two species of<br />
rice leaffolders on selected weed plants.<br />
International Rice Research Newsletter<br />
15(3): 3 1-32.<br />
Ablin, M.P. 1992. Heteropsylla sp. (Psyllidae)<br />
successfully controls pasture infestations<br />
of Mimosa invisa within three years of re-<br />
lease in Australia. p. 54, Abstracts, VIII<br />
International Symposium on Biological<br />
Control of Weeds, Lincoln University,<br />
Canterbury, New Zealand, 2-7 February<br />
1992.<br />
Ablin, M.P. 1993a. Insecticide exclusion with<br />
carbofuran demonstrates the effectiveness<br />
of Heteropsylla spinulosa as a biological<br />
control agent for Mimosa invisa in North<br />
Queensland. Abstract, 14 Asian Pacific<br />
Weed Science Society and 10th Australian<br />
Weed Society Conference, September<br />
1993, Brisbane.<br />
Ablin, M. 1993b. Successful establishment of<br />
Heteropsylla spinulosa on Mimosa invisa<br />
in Papua New Guinea. Queensland<br />
Department of Lands, Memorandum, lp.<br />
Acosta, O., Veitia, A. and Wade, J. 1986.<br />
Malezas hospedantes de Meloidogyne spp.<br />
en areas tabacaleras de la Preoincia de<br />
Sancti Spiritus. Ciencia y Tecnica en la<br />
Agricultura, Proteccion de Plantas 9(4)<br />
3 1-40. (Helminthological Abstracts,<br />
Series B, 58: 1410,1989).<br />
Adams, A. 1967. The vectors and alternate<br />
hosts of groundnut rosette virus in Central<br />
Province, Malawi. Rhodesian, Zambian,<br />
Malawian Journal of Agricultural<br />
Research 5: 145-1 5 1. (Holm et al. 1977).<br />
Agarwal, B.D. 1985. Biology of Hypolixus<br />
truncatulus Fabr. (Coleoptera:<br />
Curculionidae) forming galls on the stem<br />
of Amaranthus spinosus Linn. in India.<br />
Cecidologia Internationale 6(
234 Biological Control of Weeds: Southeast Asian Prospects<br />
Almeida, A.M.R. 1979. Experimental trans-<br />
mission of soybean mosaic virus by an<br />
aphid Dactynotus sp. occuring on Bidens<br />
pilosa, Paranh State, Brazil (Portuguese).<br />
Fitopathologia Brasiliera 4: 509-5 10.<br />
Ambika, S.R. and Jayachandra 1990. The<br />
problem of Chromolaena weed.<br />
Chromolaena odorata Newsletter 3: 1-6.<br />
Anahosur, K.H. and Sivanesan, A. 1978.<br />
Phyllachora sacchari causes tar spot or<br />
black spot on leaves of Polytoca macro-<br />
phylla, Rottboellia exaltata, Saccharum,<br />
Sorghum. Descriptions of Pathological<br />
Fungi and Bacteria, Commonwealth<br />
Mycological Institute, Kew, UK. 59(588)<br />
~PP.<br />
Ananthakrishnan, T.N. and Thangavelu, K.<br />
1976. The cereal thrips (Haplothrips gan-<br />
glbaueri Schmutz) with special reference<br />
to the trends of infestation on Oryza sativa<br />
and the weed Echinochloa crus-galli.<br />
Proceedings of the Indian Academy of<br />
Science B 83: 196-201.<br />
Ang, O.C., Poh, T.W. and Chuan, L.Y. 1977.<br />
A whitefly-borne virus disease of<br />
Ageratum conyzoides (Linn.) in Malaysia.<br />
MARDI Research Bulletin 5: 148-152.<br />
Anon 1960. Index of plant diseases in the<br />
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Agricultural Research, Canberra. 138 pp.<br />
Waterhouse, D.F. and Norris, K.R. 1987.<br />
Biological Control: Pacific Prospects.<br />
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water weevil (Coleoptera: Curculionidae)<br />
damage to rice and aquatic weeds on aster<br />
leafhopper (Homoptera: Cicadellidae) density.<br />
Environmental Entomology<br />
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purslane sawfly-Schizocerus zabriskei<br />
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Scamurius sp. (Hemiptera, Heteroptera:<br />
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15~~.<br />
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274 Biological Control of Weeds: Southeast Asian Prospects<br />
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College, Pennsylvania. 41 Opp.
6 Index of scientific names of insects<br />
abdominalis,<br />
Microcephalothrips<br />
Acanthoscelides pigricola<br />
Kingsolver Col.:<br />
Bruchidae 152<br />
Acanthoscelides puniceus<br />
Johnson Col.: Bruchidae<br />
150,152,153<br />
Acanthoscelides<br />
quadridentatus (Schaeffer)<br />
Col.: Bruchidae 150, 152,<br />
153<br />
Acanthoscelides zebratus<br />
Kingsolver Col.:<br />
Bruchidae 152<br />
Achaea janata (Linnaeus)<br />
Lep.: Noctuidae 105<br />
Acigona infusella, see<br />
Haimbachia infusella<br />
Acrida turrita Linnaeus Ort.:<br />
Acrididae 133<br />
Acrosternum herbidum (Stiil)<br />
Hem.: Pentatomidae 144<br />
Actinote anteas (Doubleday<br />
and Hewitson) Lep.:<br />
Nymphalidae 39,44,47,<br />
48<br />
aculeatum, Coelocephalapion<br />
acuminata, Paulinia<br />
acuta, Leptocorisa<br />
Adaina bipuncta Moschler<br />
Lep.: Pterophoridae 129<br />
adipalis, Ategumia<br />
aeneicollis, Horismenus<br />
aerea, Colaspis<br />
aerea, Maecolaspis<br />
Aerenica hirticornis (Klug)<br />
Col.: Cerambycidae 40<br />
afinis, Neohydronomus<br />
agamemnon, Chi10<br />
Agathis Hyrn.: Braconidae<br />
206<br />
Agraulis vanillae (Linnaeus)<br />
Lep.: Nymphalidae 187,<br />
188<br />
Agrmyza parvicornis Loew<br />
Dip.: Agromyzidae 64<br />
Agromyza proxima Spencer<br />
Dip.: Agromyzidae 65<br />
Agrotera basinotata<br />
Harnpson Lep.:<br />
Pyralidae 122<br />
Agrotis ipsilon (Hufnagel)<br />
Lep.: Noctuidae 21 2<br />
albicinctus, Orosius<br />
albiguttalis, Sameodes<br />
Alcidodes Col.: Curculionidae<br />
1 22<br />
aliphera, Eueides<br />
allecta, Calycomyza<br />
Altica cyanea (Weber) Col.:<br />
Chrysomelidae 121, 1 22<br />
Amata huebneri Boisduval<br />
Lep.: Amatidae 134<br />
Amauromyza Dip.:<br />
Agromyzidae 32,57<br />
Amauromyza maculosa<br />
(Malloch) Dip.:<br />
Agromyzidae 29,30<br />
amazonus, Longitarsus<br />
arnbrosiae, Dactynotus<br />
ambrosiae, Uroleucon<br />
Ammalo arravica, see<br />
Pareuchaetes<br />
pseudoinsulata<br />
Ammalo insulata, see<br />
Pareuchaetes insulata<br />
Amsacta gangis, see<br />
Creatonotos gangis<br />
Amsacta lactinea (Cramer)<br />
Lep.: Arctiidae 45<br />
anaphalidis, Masonaphis<br />
anaphalidis, Neomasonaphis<br />
andropogonis, Dyodiplosis<br />
andropogonis, Orseolia<br />
anisodactylus, Sphenarches<br />
Anoecia querci , see<br />
Stegophylla querci<br />
antem, Actinote<br />
antica, Pochazia<br />
antonii, Helopeltis<br />
Apanteles Hyrn.: Braconidae<br />
206<br />
Apanteles cordoi De Santis<br />
Hyrn.: Braconidae 80<br />
Apanteles creatonoti Viereck<br />
Hyrn.: Braconidae 43,<br />
Apantelesfluitantis De Santis<br />
Hyrn.: Braconidae 80<br />
Aphis Hem.: Aphididae 121<br />
Aphis citricola, see Aphis<br />
spiraecola<br />
Aphis coreopsidis (Thomas)<br />
Hem.: Aphididae 30<br />
Aphis craccivora Koch Hem.:<br />
Aphididae 13,45, 105<br />
Aphis fabae Scopoli Hem.:<br />
Aphididae 41,64, 188<br />
Aphis gossypii Glover Hem.:<br />
Aphididae 13,45, 105, 133<br />
188<br />
Aphis illinoisensis Shimer<br />
Hem.: Aphididae 30<br />
Aphis nigricauda, see Aphis<br />
spiraecola<br />
Aphis spiraecola Patch Hem.:<br />
Aphididae 1 3,41,44,45,<br />
133,188<br />
aphodoides, Temnodachrys<br />
Apion Col.: Apionidae 130,<br />
213,215<br />
Apion brunneonigrum<br />
BCguin-Billecocq Col.:<br />
Apionidae 38,39,42-44,<br />
46-48<br />
Apion luteirostre Gerstaecker<br />
Col.: Apionidae 30, 127,<br />
129,130<br />
Apsilops sp. Hyrn.:<br />
Ichneumonidae 206<br />
archboldi, Liriomyza<br />
Archips Lep.: Tortricidae<br />
23<br />
Archips micaceana (Walker)<br />
Meyrick Lep.: Tortricidae<br />
123<br />
arctithorax, Baris<br />
argentiniensis, Disonycha<br />
Argentinorhynchus bennetti<br />
O'Brien and Wibmer Col.:<br />
Curculionidae 200
278 Biological Control ( ~f Weeds: Southeast Asian Prc<br />
Argentinorhynchus breyeri<br />
Br2thes Col.:<br />
Curculionidae 200<br />
Argentinorhynchus bruchi<br />
(Hustache) Col.:<br />
Curculionidae 200,204<br />
Argentinorhynchus minimus<br />
O'Brien and Wibmer Col.:<br />
Curculionidae 200<br />
Argentinorhynchus<br />
squamosus (Hustache)<br />
Col.: Curculionidae 200<br />
argentinus, Horciacinus<br />
Argyractis subornata<br />
(Hampson) Lep.: Pyralidae<br />
72,201<br />
Argyresthia leuculias<br />
Meyrick Lep.:<br />
Argyresthiidae 122<br />
armatus, Chariesterus<br />
armigera, Dicladispa<br />
Aroga Lep.: Gelechiidae 15 1<br />
arravaca, Ammalo<br />
arravaca, Pareuchaetes<br />
artemisia, Dynamine<br />
Arthrocnodax meridionalis<br />
Felt Dip.: Cecidomyiidae<br />
48<br />
Ascia buniae phaloe, see<br />
Perrhybris phaloe<br />
aspersa, Sibinia<br />
asperulus, Ceutorhynchus<br />
Asphondylia Dip.:<br />
Cecidomyiidae 2 1 6<br />
Asphondylia bidens<br />
Johannsen Dip.:<br />
Cecidomyiidae 3 1<br />
Asphondylia corbulae Mohn<br />
Dip.: Cecidomyiidae 40<br />
Asphondylia portulacae<br />
Mohn Dip.: Cecidomyiidae<br />
213,216<br />
Astylus lineatus (Fabricius)<br />
Col.: Melyridae 22<br />
Asynonychus godmani Crotch<br />
Col.: Curculionidae 144<br />
Ategumia adipalis (Zeller)<br />
Lep.: Pyralidae 120, 122,<br />
123<br />
Ategumia fatualis (Lederer)<br />
Lep.: Pyralidae 120, 122,<br />
123<br />
Atherigona soccata Rondani<br />
Dip.: Muscidae 225<br />
atilus, Sphaeniscus<br />
atricornis, Phytomyza<br />
atrovenosa, Nisia<br />
Atractomorpha crenulata<br />
(Fabricius) Ort.: Acrididae<br />
78<br />
Aulacorthum solani<br />
(Kaltenbach) Hem.<br />
Aphididae 13<br />
Aulacochlamys Col.<br />
Chrysomelidae 39,49<br />
aurata aurantior,<br />
Pareuchaetes<br />
aurata aurata, Pareuchaetes<br />
aurata, Pareuchaetes<br />
Austelasmus Hym.:<br />
Elasmidae 5 1<br />
Autoba versicolor (Walker)<br />
Lep.: Noctuidae 122<br />
avenae, Sitobion<br />
Azarma densa, see Bellura<br />
densa<br />
azia, Thecla<br />
azia, Tmolus<br />
Bactrocera dorsalis (Hendel)<br />
Dip.: Tephritidae 120, 122<br />
Bactrocera pedestris, see<br />
Bactrocera dorsalis<br />
bada, Parnara<br />
Baliothrips biformis, see<br />
Stenchaetothrips biformis<br />
baridiiformis,<br />
Psuedoderelomus<br />
Baris Col.: Curculionidae 29,<br />
30,40<br />
Baris arctithorax (Pic) Col.:<br />
Curculionidae 2 12,213,<br />
216<br />
Baris lorata Marshall Col.:<br />
Curculionidae 2 1 3<br />
Baris portulacae Marshall<br />
Col.: Curculionidae 2 13<br />
basalis, Xyonysius<br />
basinotata, Agrotera<br />
basui, Tetraneura<br />
Bellura densa (Walker) Lep.:<br />
Noctuidae 73,79,80<br />
Bemisia costa-limai, see<br />
Bemisia tabaci<br />
Bemisia tabaci (Gennadius)<br />
Hem.: Aleyrodidae 14, 16,<br />
99,100,105<br />
bennetri, Argentinorhynchus<br />
bertrandi, Hypurus<br />
bessus, Nisoniades<br />
bibiana, Syphrea<br />
biblis, Didonis<br />
bicincta fraterna,<br />
Monecphora<br />
bicincta fraterna, Prosapia<br />
bicolor, Stirellus<br />
bidens, Asphondylia<br />
bidentis, Melanogromyza<br />
bifidalis, Loxostege<br />
biformis, Baliothrips<br />
biformis, Stenchaetothrips<br />
bigaria, Desmogramma<br />
binaria, Micrutalis<br />
bipartista, Dactylispa<br />
bipuncta, Adaina<br />
bissellata, Coelophora<br />
Blissus leucopterus (Say)<br />
Hem.: Lygaeidae 90<br />
botanephaga, Sesamia<br />
Bothrogonia ferrugenea<br />
(Fabricius) Hem.:<br />
Cicadellidae 133<br />
Brachycaudus helichrysi<br />
(Kaltenbach) Hem.:<br />
Aphididae 13<br />
Brachycyttarus griseus De<br />
Joannis Lep.: Psychidae<br />
181,182<br />
Brachymeria ovata (Say)<br />
Hym.: Chalcididae 187<br />
Brachymeria russelli Burks<br />
Hym.: Chalcididae 130<br />
breyeri, Argentinorhynchus<br />
brickelliae, Neolasioptera<br />
bruchi, Argentinorhynchus<br />
bruchi, Neochetina<br />
bruchi, Ochetina<br />
brunella, Selca<br />
brunnea, Myrmicaria<br />
brunneonigrum, Apion<br />
bryoniae, Liriomyza<br />
Bucculatrix sp. Lep.:<br />
Bucculatricidae 39,49<br />
buniae phaloe, Ascia
Caicella calchas, see Cogia<br />
calchas<br />
calamistis, Sesamia<br />
calchas, Caicella<br />
calchas, Cogia<br />
Calephelis Lep.: Lycaenidae<br />
162<br />
Calephelis laverna Godman<br />
and Salvin Lep.:<br />
Lycaenidae 40<br />
Caliothrips ipomoeae<br />
Moulton Thy.: Thripidae<br />
14<br />
Calycomyza Dip.:<br />
Agromyzidae 14,32<br />
Calycomyza allecta<br />
(Melander) Dip.:<br />
Agromyzidae 29,3 1<br />
Calycomyza mikaniae<br />
Spencer Dip.:<br />
Agromyzidae 128<br />
Calycomyza platyptera<br />
(Thornson) Dip.:<br />
Agromyzidae 29,3 1<br />
campanulicollis, Promecops<br />
Capitophorus hippophaes<br />
(Walker) Hem.: Aphididae<br />
13<br />
carduana,Lobesia<br />
carduana, Polychrosis<br />
carduellinus, Hyperomyzus<br />
cariniventris, Rhodobaenus<br />
Cannenta mimosa Eichlin<br />
and Passoa Lep.: Sesiidae<br />
147,150,152-154,156<br />
Caryedes pickeli (Pic) Col.:<br />
Bruchidae 101<br />
Cassida exilis Col.:<br />
Chrysomelidae 22<br />
Cassida nigriventris Col.:<br />
Chrysomelidae 21,22<br />
Catantops humilis Serville<br />
Ort.: Acrididae 133<br />
catenulata, Lophocampa<br />
caulophaga, Liriomyza<br />
Cecidochares fluminensis<br />
(Lima) Dip.: Tephritidae<br />
40<br />
Cecidomyia Dip.:<br />
Cecidomyiidae 193<br />
6 Index of scientific names of insects 279<br />
Cecidomyia penniseti Felt<br />
Dip.: Cecidomyiidae 193,<br />
194<br />
Centrotypus flexuosus<br />
(Fabricius) Hem.:<br />
Membracidae 133<br />
Centrinaspis Col.:<br />
Curculionidae 29,30,40<br />
Centrinaspis perscitus see<br />
Linogeraeus perscitus<br />
Ceresa ustulata Fairmaire<br />
Hem.: Membracidae 143<br />
Cerodontha muscina<br />
(Meigen) Dip.:<br />
Agromyzidae 65<br />
cervinus, Pantomorus<br />
Ceutorhynchus Col.:<br />
Curculionidae 122,215<br />
Ceutorhynchus asperulus<br />
Faust Col.: Curculionidae<br />
22<br />
Ceutorhynchus oleracae<br />
Marshall Col.:<br />
Curculionidae 2 1 3<br />
Ceutorhynchus portulacae<br />
Marshall Col.:<br />
Curculionidae 2 1 1,213<br />
Chaetogeoica graminiphaga<br />
Raychaudhuri, Pal and<br />
Ghosh Hem.: Aphididae<br />
0 0<br />
00<br />
Chalcodermus Col.:<br />
Curculionidae 144, 162<br />
Chalcodermus segnis Fielder<br />
Col.: Curculionidae 144<br />
Chalcodermus serripes<br />
Fahraeus Col.:<br />
Curculionidae 15 1, 154<br />
Chalcophana viridipennis<br />
Germar Col.:<br />
Chrysomelidae 30<br />
Chariesterus armatus<br />
(Thunberg) Hem.:<br />
Coreidae 101<br />
charitonia, Heliconius<br />
Chilo Lep.: Pyralidae 225<br />
Chilo agamemnon Bleszynski<br />
Lep.: Pyralidae 65<br />
Chilo zacconius Bleszynski<br />
Lep.: Pyralidae 65<br />
Chlamisus Col.:<br />
Chrysomelidae 16 1<br />
Chlamisus insularis (Jacoby)<br />
Col.: Chrysomelidae 49<br />
Chlamisus mimosae Karren<br />
Col.: Chrysomelidae 30,<br />
39,49,150,152,154<br />
Chloropteryx Lep.:<br />
Geometridae 129<br />
Chnootriba similis<br />
(Thunberg) Col.:<br />
Coccinellidae 225<br />
Chortogonus trachypterus<br />
Blanchard Ort.: Acrididae<br />
105<br />
Chromatomyia Dip.<br />
Agromyzidae 32<br />
Chrysolina Col.:<br />
Chrysomelidae 130<br />
Cicadulina triangula Hem.:<br />
Cicadellidae 93<br />
cinerea, Piesma<br />
Circulifer haematoceps<br />
(Mulsant and Rey) Hem.:<br />
Cicadellidae 21 1<br />
citri, Planococcus<br />
citricola, Aphis<br />
civiloides, Exorista<br />
Cladochaeta nebulosa<br />
Coquillett Dip.:<br />
Drosophilidae 3 1<br />
claripennis, Phorocera<br />
claudina, Tegosa<br />
Cleitodiplosis graminis<br />
(Tavares) Dip.:<br />
Cecidomyiidae 1 8 1, 182<br />
Cletusfuscescens Walker<br />
Hem.: Coreidae 21<br />
Clinodiplosis Dip.:<br />
Cecidomyiidae 39, 40,49<br />
Clinodiplosis eupatorii (Felt)<br />
Dip.: Cecidomyiidae 40<br />
Clovia conifer (Walker)<br />
Hem.: Cercopidae 133<br />
Cnaphalocrocis medinalis<br />
(GuenCe) Lep.: Pyralidae<br />
65,90,176,182,194<br />
Cnaphalocrocis patnalis<br />
Bradley Lep.: Pyralidae<br />
65,90,176,182<br />
Coccus Hem.: Coccidae 100
280 Biological Control of Weeds: Southeast Asian Pra<br />
Coccus longulus (Douglas)<br />
Hem.: Coccidae 161<br />
Coelocephalapion aculeatum<br />
(Fall) Col.: Apionidae 150,<br />
152,154<br />
Coelocephalapion pigrae<br />
Kissinger Col.: Apionidae<br />
151,154<br />
Coelocephalapion<br />
spretissimum (Sharp) Col.:<br />
Apionidae 154<br />
Coelophora bissellata<br />
Mulsant Col.:<br />
Coccinellidae 134<br />
Coelosternus notaticeps, see<br />
Sternocoelus notaticeps<br />
coffearia, Homona<br />
Cogia calchas (Herrich-<br />
Schaffer) Lep.:<br />
Hesperiidae 145, 162<br />
Colaspis Col.: Chrysomelidae<br />
144,181<br />
Colaspis aerea Lefevre Col.:<br />
Chysomelidae 182<br />
Coleophora versurella Zeller<br />
Lep.: Coleophoridae 21,22<br />
commelinae, Liriomyza<br />
complicata, Exema<br />
compositae, Uroleucon<br />
concinnus, Euscyrtus<br />
conifer, Clovia<br />
conjuncta, Desmogramma<br />
connexa, Eriopis<br />
connexa, Procecidochares<br />
consimilis, Phaedon<br />
Contarinia Dip.:<br />
Cecidomyiidae 40,88,93,<br />
194<br />
Contarinia panici (Plotnikov)<br />
Dip.: Cecidomyiidae 175,<br />
177<br />
Contarinia sorghicola<br />
(Coquillett) Dipt.:<br />
Cecidomyiidae 93, 193,<br />
194<br />
corbulae, Asphondylia<br />
cordoi, Apanteles<br />
coreopsidis, Aphis<br />
Corythuca pellucidu Drake<br />
and Hambleton Hem.:<br />
Tingidae 101<br />
Corythuca socia Monte<br />
Hem.: Tingidae 101<br />
costa-limai, Bemisia<br />
Courteia graminis, see<br />
Orseolia graminis<br />
craccivora, Aphis<br />
creatonoti, Apanteles<br />
Creatonotos gangis<br />
(Linnaeus) Lep.: Arctiidae<br />
90,111,182<br />
crenulata, Atractomorpha<br />
cretatus, Onychylis<br />
cretatus, Pistiacola<br />
cretica, Sesamia<br />
cribarius, Euschistus<br />
Criotettrix Ort.: Tetrigidae<br />
200,202<br />
Cropia minthe (Dyar) Lep.:<br />
Noctuidae 3 1<br />
cruttwellae, Neolasioptera<br />
Cryptocephalus miserabilis<br />
Suffrian Col.:<br />
Chrysomelidae 15 1<br />
Cryptocephalus viridiaeneus<br />
Boheman Col.:<br />
Chrysomelidae 1 44<br />
Cryptorhynchus Col.:<br />
Curculionidae 122<br />
cubana, Heteropsylla<br />
curvifascia, Erastroides<br />
cyanea, Altica<br />
cyanea, Haltica<br />
Cyanotricha necyria (Felder<br />
and Rogenhofer) Lep.:<br />
Dioptidae 188<br />
cydno, Heliconius<br />
cymoides, Nysius<br />
cynodontis. Orseolia<br />
Cyrtorhinus lividipennis<br />
Reuter Hem.: Miridae 64<br />
Dactylispa bipartista Col.:<br />
Chrysomelidae 134<br />
Dactynotus ambrosiae, see<br />
Uroleucon ambrosiae<br />
Dactynotus, see Uroleucon<br />
Delphacodes idonea Hem.:<br />
Delphacidae 176<br />
densa, Bellura<br />
depunctalis, Nymphula<br />
Desmogramma 130<br />
Desmogramma bigaria<br />
Erichson Col.:<br />
Chrysomelidae 130<br />
Desmogramma conjuncta<br />
BechynC Col.:<br />
Chrysomelidae 127, 130<br />
Diacamma rugosum (Le<br />
Guillou) Hym.: Formicidae<br />
43<br />
Diachrysia orichalcea, see<br />
Thysanoplusia orichalcea<br />
Dialeurodes vulgaris Singh<br />
Hem.: Aleyrodidae 30<br />
Dichelops fircatus<br />
(Fabricius) Hem.:<br />
Pentatomidae 144<br />
Dichomeris Lep.: Gelechiidae<br />
14,40<br />
Dicladispa armigera<br />
(Olivier) Col.:<br />
Chrysomelidae 64<br />
Didonis biblis (Fabricius)<br />
Lep.: Nymphalidae 101<br />
diemenalis, Lamprosema<br />
dilatatum, Sipylus<br />
diminutalis, Nymphula<br />
diminutalis, Paraponyx<br />
Dinarmus Hym.:<br />
Pteromalidae 152<br />
Dione juno (Cramer) Lep.:<br />
Nymphalidae 186, 1 87<br />
Dioxyna picciola, see<br />
Dioxyna sororcula<br />
Dioxyna sororcula<br />
(Wiedemann) Dip.:<br />
Tephritidae 3 1<br />
Diplacaspis miserabilis, see<br />
Cryptocephalus<br />
miserabilis<br />
Diplacaspis prosternalis<br />
(Schaeffer) Col.:<br />
Chrysomelidae 15 1<br />
Disonycha argentiniensis<br />
BechynC Col.:<br />
Chrysomelidae 10 1<br />
distincta, Lamprosema<br />
dolosa, Pegomya<br />
dorsalis, Bactrocera<br />
dorsalis, Recilia<br />
drumalis, Petrophila<br />
Dryas julia (Fabricius) Lep.:<br />
Nymphalidae 187
Dynamine artemisia<br />
(Fabricius) Lep.:<br />
Nymphalidae 101<br />
Dyodiplosis andropogonis,<br />
see Orseolia andropogonis<br />
Dyodiplosisfluitans, see<br />
Lasioptera fluitans<br />
Dyodiplosisfluvialis, see<br />
Orseolia fluvialis<br />
Dyops minthe, see Cropia<br />
minthe<br />
echeclus, Hippotion<br />
echinochloa, Lasioptera<br />
Echinocnemus Col.:<br />
Curculionidae 1 15<br />
Echoma marginata<br />
(Linnaeus) Col.:<br />
Chrysomelidae 127, 1 30<br />
Echoma quadristillata<br />
(Boheman) Col.:<br />
Chrysomelidae 127, 130<br />
Ecpantheria hambletoni, see<br />
Hypercompe hambletoni<br />
Edessa meditabunda<br />
(Fabricius) Hem.:<br />
Pentatomidae 144<br />
eichhorniae, Neochetina<br />
Elachertus Hym.: Eulophidae<br />
46<br />
Elasmopalpus lignosellus<br />
(Zeller) Lep.: Pyralidae 65<br />
Elasmus Hym.: Elasmidae 46<br />
Eldana saccharins Walker<br />
Lep.: Pyralidae 65<br />
elegans, Neohydronomus<br />
elimatus, Dalbulus<br />
Elophila responsalis<br />
(Walker) Lep.: Pyralidae<br />
115,116,167,201,202<br />
ementitus, Xyonysius<br />
Emmalocera Lep.: Pyralidae<br />
63,65<br />
Enchenopa gracilis (Germar)<br />
Hem.: Membracidae 143<br />
entreriana, Tomaspis<br />
entreriana, Zulia<br />
Entylia Hem.: Membracidae<br />
128<br />
Entylia sinuata (Fabricius)<br />
Hem.: Membracidae 128<br />
6 Index of scientific names of insects 281<br />
Epilachna indica Mulsant<br />
Col.: Coccinellidae 134<br />
Epilachna similis, see<br />
Chnootriba similis<br />
Epipagis albiguttalis, see<br />
Sameodes albiguttalis<br />
Epipsammia pectinicornis,<br />
see Spodoptera<br />
pectinicornis<br />
Episammea pectinicornis, see<br />
Epipsammia pectinicornis<br />
Episcada pascua Schaus<br />
Lep.: Nymphalidae 101<br />
Erastroides curvifascia<br />
Hampson Lep.: Noctuidae<br />
200<br />
erato, Heliconius<br />
Eretmocera impactella<br />
(Walker) Lep.:<br />
Scythrididae 23<br />
eridania, Prodenia<br />
eridania, Spodoptera<br />
Erinnyis oenotrus (Cramer)<br />
Lep.: Sphingidae 101<br />
Eriopis connexa (Germar)<br />
Col.: Coccinellidae 141<br />
Eryphanis polyxena<br />
Meerburgh Lep.:<br />
Nymphalidae 193, 194<br />
erythrocephalus, Stiretrus<br />
etheiella, Eucampyla<br />
Eublemma versicolor, see<br />
Autoba versicolor<br />
Eucalymnatus Hem.:<br />
Coccidae 100<br />
Eucampyla etheiella Meyrick<br />
Lep.: Pyralidae 46,47<br />
Euderus Hym.: Eulophidae<br />
52<br />
Eueides aliphera Godart<br />
Lep.: Nymphalidae 187<br />
Eueides isabella (Cramer)<br />
Lep.: Nymphalidae 187<br />
Eueides procula Butler and<br />
Drull Lep.: Nymphalidae<br />
187<br />
eupatoriella, Melanagromyza<br />
eupatorii, Clinodiplosis<br />
eupatorii, Neolasioptera<br />
euphorbiae, Haplothrips<br />
Eupithecia Lep.: Geometridae<br />
129<br />
Eurema Lep.: Pieridae 139<br />
Eurema lisa (Boisduval and<br />
LeConte) Lep.: Pieridae<br />
162<br />
Eurema tenella (Boisduval)<br />
Lep.: Pieridae 145, 162<br />
Euschistus tristigmus<br />
cribarius Stdl Hem.:<br />
Pentatomidae 144<br />
Euschistus luridus Dallas<br />
Hem.: Pentatomidae 144<br />
Euscyrtus concinnus (de<br />
Haan) Ort.: Gryllidae 57,<br />
64,224<br />
Exema complicata Jacoby<br />
Col.: Chrysomelidae 128,<br />
132<br />
exigua, Spodoptera<br />
exilis, Cassida<br />
Exorista Dip.: Tachinidae 43<br />
Exorista civiloides, see<br />
Exorista xanthaspis<br />
Exorista xanthaspis<br />
Wiedemam Dip.:<br />
Tachinidae 45<br />
explanata, Pentispa<br />
fabae, Aphis<br />
fasciatus, Pistiacola<br />
fascifrons, Macrosteles<br />
fastigiata, Sibinia<br />
fatualis, Ategumia<br />
femorata, Sagra<br />
Ferrisia virgata (Cockerell)<br />
Hem.: Pseudococcidae 105<br />
ferrugenea, Bothrogonia<br />
ferruginae, Temlucha<br />
flexuosus, Centrotypus<br />
floris, Melanagromyza<br />
fluitans, Dyodiplosis<br />
fluitans, Lasioptera<br />
fluitantis, Apanteles<br />
fluminensis, Cecidochares<br />
fluvialis, Dyodiplosis<br />
fluviatilis, Orseolia<br />
fregonalis, Nymphula<br />
frugiperda, Spodoptera<br />
frugivora, Neolasioptera<br />
furcatus, Dichelops
282 Biological Control of Weeds: Southeast Asian Prospects<br />
furcifera, Sogatella<br />
furnacalis, Ostrinia<br />
fuscesens, Cletus<br />
gangis, Amsacta<br />
gangis, Creatonotos<br />
ganglbaueri, Haplothrips<br />
Garcanus gracilentus (St&)<br />
Hem.: Miridae 30<br />
Gargara Hem.: Membracidae<br />
121<br />
geminata, Solenopsis<br />
gentilei, Tetrastichus<br />
gentilei, Thripsastichus<br />
Geoica Iucifuga (Zehntner)<br />
Hem.: Aphididae 88<br />
Germalus unipunctatus<br />
(Montandon) Hem.:<br />
Lygaeidae 21<br />
Geromyia penniseti (Felt)<br />
Dip.: Cecidomyiidae 193,<br />
194<br />
Geromyia seminis (Felt) Dip.:<br />
Cecidomyiidae 193,194<br />
Gesonula punctifrons (Sdl)<br />
Ort.: Acrididae 78, 167<br />
Gibbobruchus pickeli , see<br />
Caryedes pickeli<br />
Gibbobruchus polycoccus<br />
(Fahraeus) Col.: Bruchidae<br />
101<br />
godmani, Asynonychus<br />
gossypii, Aphis<br />
gowdeyi, Haplothrips<br />
gracilentus, Garcanus<br />
gracilis, Enchenopa<br />
graminiphaga, Chaetogeoica<br />
graminis, Cleitodiplosis<br />
graminis, Courteia<br />
graminis, Orseolia<br />
graminum, Schizaphis<br />
granarium, Macrosiphum<br />
griseola, Hydrellia<br />
griseus, Brachycyttarus<br />
guerini, Lexiphanes<br />
guildinii, Piezodorus<br />
gunniella, Neurostrota<br />
Gyrocnemis Hem.:<br />
Thyreocoridae 144<br />
haematoceps, Circulijier<br />
I<br />
Haimbachia infusella<br />
(Walker) Lep.: Pyralidae<br />
69,73,74,76,79,80,82<br />
Haltica cyanea, see Altica<br />
cyanea<br />
Halticus minutus Reuter<br />
Hem.: Miridae 132<br />
hambletoni, Ecpantheria<br />
hambletoni, Hypercompe<br />
hanno, Hemiargus<br />
Haplopeodes Dip.:<br />
Agromyzidae 21 6<br />
Haplopeodes minutus (Frost)<br />
Dip.: Agromyzidae 21,22<br />
Haplopeodes palliatus<br />
(Czoquillett) Dip.:<br />
Agromyzidae 2 12,2 1 3,<br />
216<br />
Haplothrips Thy.:<br />
Phlaeothripidae 134<br />
Haplothrips euphorbiae Thy.:<br />
Phlaeothripidae 105<br />
Haplothrips ganglbaueri<br />
Schmutz Thy.:<br />
Phlaeothripidae 63, 64, 89<br />
Haplothrips gowdeyi<br />
(Franklin) Thy.:<br />
Phlaeothripidae 14, 101,<br />
182<br />
Haplothrips longisetosus<br />
Ananthakrishnan Thy.:<br />
Phlaeothripidae 22<br />
Haplothrips paumalui<br />
Moulton Thy.:<br />
Phlaeothripidae 182<br />
harleyi, Teleonemia<br />
hawaiiensis, Thrips<br />
hecale, Heliconius<br />
helichrysi, Brachycaudus<br />
Heliconius charitonia<br />
(Linnaeus) Lep.:<br />
Nymphalidae 188<br />
Heliconius cydno Bates Lep.:<br />
Nymphalidae 188<br />
Heliconius erato Doubleday<br />
Lep.: Nymphalidae 188<br />
Heliconius hecale Hewitson<br />
Lep.: Nymphalidae 187,<br />
189<br />
Helicoverpa obsoleta, see<br />
Helicoverpa zea<br />
Helicoverpa virescens, see<br />
Heliothis virescens<br />
Helicoverpa zea (Boddie)<br />
Lep.: Noctuidae 188<br />
Heliodines Lep.: Heliodinidae<br />
216<br />
Heliodines quinqueguttata<br />
Walshingham Lep.:<br />
Heliodinidae 2 13,2 16<br />
Heliothis virescens<br />
(Fabricius) Lep.:<br />
Noctuidae188<br />
Hellula undalis (Fabricius)<br />
Lep.: Pyralidae 134<br />
Helopeltis Hem.: Miridae 133<br />
Helopeltis antonii Signoret<br />
Hem.: Miridae 121<br />
Hemiargus hanno (Stoll)<br />
Lep.: Lycaenidae 145, 161,<br />
162<br />
Hemiberlesia lataniae<br />
(Signoret) Hem.:<br />
Diaspididae 161<br />
hennia, Proxenus<br />
herbidum, Acrosternum<br />
Heteropsylla Hem.: Psyllidae<br />
139,142,144,161<br />
Heteropsylla cubana<br />
Craw ford Hem.: Psyllidae<br />
1 42<br />
Heteropsylla spinulosa<br />
Muddiman, Hodkinson and<br />
Hollis Hem.: Psyllidae<br />
137,139-1 45<br />
hieroglyphicus, Poekilocerus<br />
Hilar Col.: Chrysomelidae<br />
1 44<br />
Hippotion echeclus<br />
(Boisduval) Lep.:<br />
Sphingidae 78, 167<br />
hirsuta, Tetraneura<br />
hirticornis, Aerenica<br />
Homoeocerus serrijier<br />
Westwood Hem.: Coreidae<br />
133<br />
Homona cofearia (Nietner)<br />
Lep.: Tortricidae 45<br />
Horciacinus argentinus, see<br />
Horciacinus signoreti<br />
Horciacinus signoreti (Berg)<br />
Hem.: Miridae 143
Horcias nobilellus (Berg)<br />
Hem.: Miridae 22,30<br />
Horismenus ?aeneicollis<br />
Ashmead Hym.:<br />
Eulophidae 130, 132<br />
hospes, Spilostethus<br />
huebneri, Amata<br />
humilis, Catantops<br />
humilis, Hyperodes<br />
humilis, Listronotus<br />
Hyalomyodes triangulifer<br />
(Loew) Dip.: Tachinidae<br />
130<br />
Hyalopeplus vitripennis (Stll)<br />
Hem.: Miridae 121<br />
Hydrellia griseola (Falltn)<br />
Dip.: Ephydridae 65<br />
Hymenia recurvalis, see<br />
Spodoptera recurvalis<br />
Hypanthus Col.:<br />
Curculionidae 144<br />
Hypercompe hambletoni<br />
(Schaus) Lep.: Arctiidae<br />
3 1<br />
Hyperodes humilis, see<br />
Listronotus humilis<br />
Hyperomyzus carduellinus<br />
(Theobald) Hem.:<br />
Aphididae 13<br />
Hypolixus ritsemae (Pascoe)<br />
Col.: Curculionidae 22<br />
Hypolixus trunculatus<br />
(Fabricius) Col.:<br />
Curculionidae 19,21,22,<br />
45<br />
Hypomicrogaster Hym.:<br />
Braconidae 81<br />
Hypurus bertrandi Pems<br />
Col.: Curculionidae<br />
21 1-213,215-217<br />
Hysteroneura setariae<br />
(Thomas) Hem.:<br />
Aphididae 89<br />
Icerya seychellarum<br />
(Westwood) Hem.:<br />
Margarodidael61<br />
Idiophantis Lep.: Gelechiidae<br />
122<br />
idonea, Delphacodes<br />
illinoisensis, Aphis<br />
6 Index of scientific names of insects 283<br />
Imerodes Col.: Curculionidae<br />
122<br />
impactella, Eretmocera<br />
ina, Taractrocera<br />
inaequalis, Xyonysius<br />
includens, Pseudoplusia<br />
inconspicuus, Nysius<br />
incurvus, Pycnoderes<br />
indica, Epilachna<br />
indica, Mycodiplosis<br />
infusella, Acigona<br />
infusella, Haimbachia<br />
insecta, Xanthaciura<br />
insignis, Liriomyza<br />
insularis, Chlamisus<br />
insulata, Pareuchaetes<br />
insulata, Ammalo<br />
inustorum, Lasioptera<br />
ipomoeae, Caliothrips<br />
ipsilon, Agrotis<br />
irrorata. Milothris<br />
isabella, Eueides<br />
Isothrips Thy.: Thripidae 1 33<br />
Itonida penniseti, see<br />
Geromyia penniseti<br />
Itonida seminis, see<br />
Geromyia seminis<br />
janata, Achaea<br />
javanica, Parallelodiplosis<br />
julia, Dryas<br />
juno, Dione<br />
kanni, Lasioptera<br />
knysna, Zizeeria<br />
kolophon, Sogatella<br />
krupta, Zizeeria<br />
Lactica sp. Col.:<br />
Chrysomelidae 144<br />
lactinea, Amsacta<br />
laevis, Macrotracheliella<br />
Lamprosema diemenalis<br />
(Guenke) Lep.: Pyralidae<br />
132,133<br />
Lamprosema distincta (Kaye)<br />
Lep.: Pyralidae 129<br />
Laodelphax striatellus<br />
(Falltn) Hem.:<br />
Delphacidae 64,89,93<br />
laphygmae, Meteorus<br />
Lasioptera Dip.:<br />
Cecidomyiidae 18 1<br />
Lasioptera echinochloa Felt<br />
Dip.: Cecidomyiidae 63,<br />
65<br />
Lasiopterafluitans Felt Dip.:<br />
Cecidomyiidae 177<br />
Lasioptera inustorum Felt<br />
Dip.: Cecidomyiidae 177<br />
Lasioptera kanni Felt Dip.:<br />
Cecidomyiidae 177<br />
Lasioptera panici Felt Dip.:<br />
Cecidomyiidae 177<br />
Lasioptera paniculi Felt Dip.:<br />
Cecidomyiidae 177<br />
Lasioptera portulaceae Felt<br />
Dip.: Cecidomyiidae 21 3<br />
lataniae, Hemiberlesia<br />
latigenis, Notiphila<br />
latipes, Mocis<br />
latiuscula, Scotinophara<br />
laverna, Calepheles<br />
leelamaniae, Macrosiphum<br />
leelamaniae, Sitobion<br />
Leptocentrus taurus<br />
(Fabricius) Hem.:<br />
Membracidae 12 1<br />
Leptocorisa acuta (Thunberg)<br />
Hem.: Alydidae 99<br />
Leptocorisa oratorius<br />
(Fabricius) Hem.:<br />
Alydidae 64,99<br />
Leptocorisa solomonensis<br />
Ahmad Hem.: Alydidae 99<br />
Leptocysta sexnebulosa (Stll)<br />
Hem.: Tingidae 128<br />
leucaenae, Tamarixia<br />
leucopterus, Blissus<br />
leuculias, Argyresthia<br />
Lexiphanes Col.:<br />
Chrysomelidae 144<br />
Lexiphanes guerini (Perbose)<br />
Col.: Chrysomelidae 15 1<br />
Lexiphanes semicyaneus<br />
(Suffrian) Col.:<br />
Chrysomelidae 144<br />
licarsisalis, Herpetogramma<br />
Eignosellus, Elasmopalpus<br />
linearis, Riptortus<br />
lineatus, Astylus<br />
Linogeraeus perscitus<br />
(Herbst) Curculionidae 2 13
284 Biological Control of Weeds: Southeast Asian Prospects<br />
Liothrips mikaniae (Priesner)<br />
Thy.: Phlaeothripidae 125,<br />
127,130,134,135<br />
liothrips, Thripsobremia<br />
Liriomyza Dip.: Agromyzidae<br />
31,32<br />
Liriomyza archboldi Frost<br />
Dip.: Agromyzidae 29,3 1<br />
Liriomyza bryoniae<br />
(Kaltenbach) Dip.:<br />
Agromyzidae 105<br />
Liriomyza caulophaga<br />
(Kleinschmidt) Dip.:<br />
Agromyzidae 21 1,212<br />
Liriomyza commelinae Frost<br />
Dip.: Agromyzidae 57<br />
Liriomyza insignis Spencer<br />
Dip.: Agromyzidae 29,3 1<br />
Liriomyza marginalis<br />
Malloch Dip.:<br />
Agromyzidae 89<br />
Liriomyza robustae Spencer<br />
Dip.: Agromyzidae 57<br />
Liriomyza strigata Meigen<br />
Dip.: Agromyzidae 105<br />
Liriomyza trifolii (Burgess)<br />
Dip.: Agromyzidae 3 1,211<br />
Liriomyza venegasiae<br />
Spencer Dip.:<br />
Agromyzidae 29,3 1<br />
lisa, Eurema<br />
Lissorhoptrus oryzophilus<br />
Kuschel Col.:<br />
Curculionidae 64<br />
Listronotus humilis<br />
(Gyllenhall) Col.:<br />
Curculionidae 63,64<br />
litura, Spodoptera<br />
lividipennis, Cyrtorhinus<br />
Lixophaga Dip.: Tachinidae<br />
206<br />
Lobesia carduana (Busck)<br />
Lep.: Tortricidae 129<br />
longifircifera, Sogatella<br />
longisetosus, Haplothrips<br />
Longitarsus Col.:<br />
Chrysomelidae 13 1, 132<br />
Longitarsus nr amazonus<br />
Baly Col.: Chrysomelidae<br />
128,132<br />
longulus, Coccus<br />
Lophocampa catenulata<br />
(Hubner) Lep.: Arctiidae<br />
161,162<br />
lorata, Baris<br />
Loxostege Lep.: Pyralidae 23<br />
Loxostege bijidalis<br />
(Fabricius) Lep.: Pyralidae<br />
212<br />
lucifuga, Geoica<br />
lugens, Nilaparvata<br />
luridus, Euschistus<br />
luteirostre, Apion<br />
macminni, Cerodontha<br />
rnacrophthalma, Diopsis<br />
Macrosiphum granarium, see<br />
Sitobion avenue<br />
Macrosiphum leelamaniae,<br />
see Sitobion leelamaniae<br />
Macrosiphum miscanthi, see<br />
Sitobion miscanthi<br />
Macrosiphum solidaginis, see<br />
Uroleucon solidaginis<br />
Macrosteles fascifrons (Still)<br />
Hem.: Cicadellidae 167<br />
maculosa, Amauromyza<br />
Maecolaspis aerea, see<br />
Colaspis aerea<br />
maidis, Dalbulus<br />
maidis, Peregrinus<br />
malayanus, Nephotettix<br />
Mansonia Dip.: Culicidae<br />
199<br />
Marasmia patnalis, see<br />
Cnaphalocrocis patnalis<br />
marginalis, Liriomyza<br />
marginata, Echoma<br />
Masonaphis anaphalidis, see<br />
Neomasonavhis<br />
anaphalidis<br />
mauritia, Spodoptera<br />
mazans, Staphylus<br />
medinalis, Cnaphalocrocis<br />
meditabunda, Edessa<br />
Melanagromyza Dip.:<br />
Agromyzidae 32<br />
Melanagromyza bidentis<br />
Spencer Dip.:<br />
Agromyzidae 29,3 1<br />
Melanagromyza eupatoriella<br />
Spencer Dip.:<br />
Agromyzidae 39,44,45<br />
Melanagromyza floris<br />
Spencer Dip.:<br />
Agromyzidae 29,3 1<br />
Melanagromyza metallica<br />
(Thomson) Dip.:<br />
Agromyzidae 13, 14<br />
Melanagromyza polyphyta<br />
Kleinschmidt Dip.:<br />
Agromyzidae 1 88<br />
Melanagromyza splendida<br />
Frick Dip.: Agromyzidae<br />
3 1<br />
rneridionalis, Arthrocnoda.<br />
Mescinia parvula (Zeller)<br />
Lep.: Pyralidae 39,42,44,<br />
46,49,50<br />
metallica, Melanagromyza<br />
Metqonycha pallidula<br />
(Boheman) Col.:<br />
Chrysomelidae 144<br />
Meteorus Hym.: Braconidae<br />
121<br />
micaceana, Archips<br />
Microcephalothrips Thy.:<br />
Thripidae 133<br />
Microcephalothrips<br />
abdominalis (D.L.<br />
Crawford) Thy.: Thripidae<br />
14<br />
Microcomps~s, see<br />
Microporus<br />
Microporus Hem.: Cydnidae<br />
161<br />
Micrutalis Hem.:<br />
Membracidae 143<br />
Micrutalis binaria<br />
(Fairmaire) Hem.:<br />
Membracidae 128<br />
mikaniae, Calycomyza<br />
mikaniae, Liothrips<br />
Milothris irrorata (Fabricius)<br />
Col.: Cerambycidae 153<br />
mimosa, Carmenta<br />
mimosae, Chlamisus<br />
minimus, Argentinorhynchus<br />
minthe, Cropia<br />
minthe, Dyops<br />
minuta, Oxya<br />
minutispina, Nilautama<br />
minutus, Halticus<br />
minutus, Haplopeodes
misantlensis, Pareuchaetes<br />
miscanthi, Macrosiphum<br />
miscanthi, Sitobion<br />
miserabilis, Cryptocephalus<br />
miserabilis, Diplacaspis<br />
Mocis latipes (GuenCe) Lep.:<br />
Noctuidae 3 1<br />
Monecphora bicincta<br />
fraterna, see Prosapia<br />
bicincta fraterna<br />
multiplicalis, Samea<br />
multistrigata, Phalonidia<br />
muscina, Cerodontha<br />
Mycetaspis personata<br />
(Comstock) Hem.:<br />
Diaspididae 14<br />
Mycodiplosis indica Felt<br />
Dip.: Cecidomyiidae 194<br />
Myrmicaria brunnea<br />
Saunders Hym.:<br />
Formicidae 43<br />
Mythimna unipuncta<br />
(Haworth) Lep.: Noctuidae<br />
65<br />
Myzus ornatus Laing Hem.:<br />
Aphididae 13<br />
Myzus persicae (Sulzer)<br />
Hem.: Aphididae 13,21,<br />
24, 188., 21 1,224<br />
Namangana pectinicornis,<br />
see Epipsammia pectini<br />
cornis<br />
nebulosa, Cladochaeta<br />
necyria, Cyanotricha<br />
Neochetina Col.:<br />
Curculionidae 73,76,78<br />
Neochetina bruchi Hustache<br />
Col.: Curculionidae 69,73,<br />
74,76-82<br />
Neochetina eichhorniae<br />
Warner Col.:<br />
Curculionidae 69,73-80,<br />
82<br />
Neogalea sunia (GuenCe)<br />
Lep.: Noctuidae 22,212<br />
Neohydronomus afJinis<br />
Hustache Col.:<br />
Curculionidae 197, 199,<br />
200,202-206<br />
6 Index of scientific names of insects 285<br />
Neohydronomus elegans<br />
O'Brien and Wibmer Col.:<br />
Curculionidae 200<br />
Neohydronomus pulchellus<br />
Hustache Col.:<br />
Curculionidae 200,205<br />
Neolasioptera Dip.:<br />
Cecidomyiidae 128, 132,<br />
217<br />
Neolasioptera brickelliae<br />
Mohn Dip.: Cecidomyiidae<br />
40<br />
Neolasioptera cruttwellae<br />
Gagne Dip.:<br />
Cecidomyiidae 40<br />
Neolasioptera eupatorii (Felt)<br />
Dip.: Cecidomyiidae 40<br />
Neolasioptera frugivora<br />
Gagne Dip.:<br />
Cecidomyiidae 40<br />
Neolasioptera portulacae<br />
(Cook) Dip.:<br />
Cecidomyiidae 21 3,2 17<br />
Neomasonaphis anaphalidis<br />
(Basu) Hem.: Aphididae<br />
13<br />
Nephotettix Hem.:<br />
Cicadellidae 133<br />
Nephotettix malayanus<br />
Ishihara and Kawase<br />
Hem.: Cicadellidae 89<br />
Nephotettix nigropictus, see<br />
Nephotettix<br />
nigromaculatus<br />
Nephotettix nigromaculatus<br />
(Motschulsky) Hem.:<br />
Cicadellidae 89,225<br />
Nephotettix virescens<br />
(Distant) Hem.:<br />
Cicadellidae 89<br />
nervosa, Nisia<br />
Nesoclutha pallida (Evans)<br />
Hem.: Cicadellidae 93<br />
Neurostrota gunniella<br />
(Busck) Lep.:<br />
Gracillariidae 147, 150,<br />
152,154156,162<br />
nigriabdominalis, Tetraneura<br />
nigricauda, Aphis<br />
nigrifrons, Graminella<br />
nigrirostris, Pistiacola<br />
nigriventris, Cassida<br />
nigromaculatus, Nephotettix<br />
nigropictus, Nephotettix<br />
Nilaparvata lugens (StAl)<br />
Hem.: Delphacidae 64<br />
Nilautama minutispina<br />
Funkhouser Hem.:<br />
Membracidae 121<br />
Nisaga simplex Walker Lep.:<br />
Eupterotidae 176<br />
Nisia atrovenosa, see Nisia<br />
nervosa<br />
Nisia nervosa (Motschulsky)<br />
Hem.: Meenoplidae 199,<br />
200,202<br />
Nisoniades bessus, see Cogia<br />
calchas<br />
noacki, Platinglisia<br />
nobilellus, Horacias<br />
Nodonota Col.:<br />
Chrysomelidae 144<br />
notaticeps, Coelosternus<br />
notaticeps, Sternocoelus<br />
Notiphila latigenis Hendel<br />
Dip.: Ephydridae 1 15,116<br />
Notophila similis de Meijere<br />
Dip.: Ephydridae 1 15,116<br />
nymphaeae, Rhopalosiphum<br />
Nymphula depunctalis, see<br />
Parapoynx stagnalis<br />
Nymphula diminutalis<br />
Snellen, see Parapoynx<br />
diminutalis<br />
Nymphulafregonalis Snellen<br />
Lep.: Pyralidae 167<br />
Nymphula responsalis, see<br />
Elophila responsalis<br />
Nymphula tenebralis, see<br />
Parapoynx tenebralis<br />
Nymphula turbata, see<br />
Parapoynx turbata<br />
Nysius Hem.:Lygaeidae 21<br />
Nysius cymoides (Spinola)<br />
Hem.: Lygaeidae 21 1<br />
Nysius inconspicuus Distant<br />
Hem.: Lygaeidae 14, 105<br />
Nysius vinitor Bergroth<br />
Hem.: Lygaeidae 21 1<br />
obliteralis, Synclita<br />
obsoleta, Helicoverpa
286<br />
Biological Control of Weeds: Southeast Asian Prospects<br />
Ochetina bruchi Hustache<br />
Col.: Curculionidae 200<br />
ochreosa, Sibinia<br />
Odontomachus simillimus Fr.<br />
Smith Hym.: Formicidae<br />
43<br />
Oecophylla smaragdina<br />
(Fabricius) Hym.:<br />
Formicidae 43<br />
oenotrus, Erinnyis<br />
Olcella pleuralis Becker<br />
Dip.: Chloropidae 14<br />
oleracae, Ceutorhynchus<br />
Omoplata rnarginata, see<br />
Echoma marginata<br />
Onebala tegulella<br />
(Walshingham) Lep.:<br />
Gelechiidae 128<br />
. Onychylis cretatus, see<br />
Pistiacola cretatus<br />
Onychylis sp. nr nigrirostris,<br />
see Pistiacola sp. nr<br />
nigrirostris<br />
oo, Plusia<br />
oratorius, Leptocorisa<br />
orichalcea, Diachrysia<br />
orientalis, Orosius<br />
ornatus, Myzus<br />
Orosius albicinctus, see<br />
Orosius orientalis<br />
Orosius orientalis<br />
(Matsumura) Hem.:<br />
Cicadellidae 21 1<br />
Orseolia Dip.: Cecidomyiidae<br />
87<br />
Orseolia andropogonis Felt<br />
Dip.: Cecidomyiidae 177<br />
Orseolia cynohntis Kieffer<br />
and Massalongo Dip.:<br />
Cecidomyiidae 177<br />
Orseoliafluvialis Dip.:<br />
Cecidomyiidae 177<br />
Orseoliafluviatilis (Felt)<br />
Dip.: Cecidomyiidae 88,<br />
90<br />
Orseolia graminis (Kreffer<br />
and Docters Van Leeuwen-<br />
Reijnvaan) Dip.:<br />
Cecidomyiidae 177<br />
Orseolia oryzae (Wood-<br />
Mason) Dipt.:<br />
Cecidomyiidae 65,90,<br />
175,177<br />
oryzae, Orseolia<br />
oryzae, Pachydiplosis<br />
oryzivora, Thaia<br />
oryzophilus, Lissorhoptrus<br />
Ostriniafurnacalis (GuenCe)<br />
Lep.: Pyralidae 90<br />
ovata, Brachymeria<br />
Oxya minuta Carl Ort.:<br />
Acrididae 78<br />
Oxysychus Hym.:<br />
Pteromalidae 21<br />
Pachybrachys Col.:<br />
Chrysomelidae 144<br />
Pachydiplosis, see Orseolia<br />
SP-<br />
Pachydiplosis oryzae, see<br />
Orseolia oryzae<br />
pacificus, Brachyplatys<br />
pacificus, Planococcus<br />
palegon, Thecla<br />
palegon, Thereus<br />
palliata, Phytomyza<br />
palliatus, Haplopeodes<br />
pallida, Nesoclutha<br />
pallidula, Metaxyonycha<br />
pallidulus, Taylorilygus<br />
panici, Contarinia<br />
panici, Lasioptera<br />
panici, Stenodiplosis<br />
panicola, Sogatella<br />
paniculi, Lasioptera<br />
Pantomorus cervinus, see<br />
Asynonychus godrnani<br />
Parallelodiplosis Dip.:<br />
Cecidomyiidae 177<br />
Parallelodiplosis javanica<br />
Felt Dip.: Cecidomyiidae<br />
177<br />
Parallelodiplosis paspali<br />
Dip.: Cecidomyiidae 18 1<br />
Parapoynx diminutalis<br />
Snellen Lep.: Pyralidae<br />
20 1<br />
Parapoynx stagnalis (Zeller)<br />
Lep.: Pyralidae 176, 182<br />
Parapoynx tenebralis<br />
(Lower) Lep.: Pyralidae<br />
199,20 1<br />
Parapoynx turbata Butler<br />
Lep.: Pyralidae 199,201<br />
Pareuchaetes Lep.: Arctiidae<br />
38,40<br />
Pareuchaetes arravaca<br />
(Jordan) Lep.: Arctiidae 40<br />
Pareuchaetes aurata<br />
aurantior Rothschild Lep.:<br />
Arctiidae 40<br />
Pareuchaetes aurata aurata<br />
(Butler) Lep.: Arctiidae 40,<br />
42,47,50<br />
Pareuchaetes insulata<br />
(Walker) Lep.: Arctiidae<br />
40<br />
Pareuchaetes misantlensis<br />
Rego Barros Lep.:<br />
Arctiidae 40<br />
Pareuchaetes pseuhinsulata<br />
Rego Barros Lep.:<br />
Arctiidae 14,35,38,39,<br />
42-47,50,52<br />
Parnara bada (Moore) Lep.:<br />
Hesperiidae 194<br />
Parthenothrips Thy.:<br />
Thripidae 134<br />
parvicornis, Agromyza<br />
parvula, Mescinia<br />
parvus, Phenacoccus<br />
pascua, Episcada<br />
paspali, Parallelodiplosis<br />
patnalis, Cnaphalocrocis<br />
patnalis, Marasmia<br />
Paulinia acuminata De Geer<br />
Ort.: Acrididae 200<br />
paumalui, Haplothrips<br />
pectinicornis, Epipsammia<br />
pectinicornis, Spohptera<br />
pedestris, Bactrocera<br />
Pegomya dolosa Stein Dip.:<br />
Anthomyiidae 21 3,217<br />
pellucida, Corythuca<br />
pennatula, Psalis<br />
penniseti, Cecidomyia<br />
penniseti, Geromyia<br />
penniseti, ltonida<br />
penniseti, Sesamia<br />
Pentispa explanata (Chapuis)<br />
Col.: Chrysomelidae 39,<br />
5 1<br />
Perasphondylia reticulata<br />
Mohn Dip.: Cecidomyiidae<br />
39,51
perditor, Thyanta<br />
Peregrinus maidis (Ashmead)<br />
Hem.: Delphacidae 89,225<br />
perlata, Ptychamalia<br />
Perrhybris phaloe (Godart)<br />
Lep.: Pieridae 3 1<br />
perscitus, Centrinaspis<br />
perscitus, Linogeraeus<br />
persicae, Myzus<br />
personata, Micetaspis<br />
pertinax, Phaedon<br />
peruana, Sibinia<br />
Petrophila drumalis (Dyar)<br />
Lep.: Pyralidae 201<br />
Phaedon consimilis, see<br />
Phaedon pertinax<br />
Phaedon pertinax StAl Col.:<br />
Chrysomelidae 29,30<br />
phaloe, Perrhybris<br />
Phalonidia multistrigata<br />
Walshingham Lep.:<br />
Tortricidae 129<br />
philippinensis,<br />
Pseudonapomyza<br />
Pholetesor sp.<br />
(circumscriptus group)<br />
Hym.: Braconidae 2 1 6<br />
Phorocera claripennis Pales<br />
Dip.: Tachinidae 187<br />
Physimerus Col.:<br />
Chrysomelidae 13 1<br />
Physimerus pygmaeus Jacoby<br />
Col.: Chrysomelidae 30,<br />
127,131<br />
Phytomyza atricornis Meigen<br />
Dip.: Agromyzidae 3 1<br />
Phytomyza palliata, see<br />
Haplopeodes palliatus<br />
picciola, Dioxyna<br />
pickeli, Caryedes<br />
pickeli, Gibbobruchus<br />
pieridis, Vesiculaphis<br />
Piesma cinereum Say Hem.:<br />
Piesmatidae 22<br />
Piezodorus guildinii<br />
(Westwood) Hem.:<br />
Pentatomidae 144<br />
pigrae, Coelocephalapion<br />
pigricola, Acanthoscelides<br />
pilicornis, Schizocerella<br />
6 Index of scientific names of insects 287<br />
Pinnaspis strachani (Cooley)<br />
Hem.: Diaspididae 16 1<br />
Pionea upalusalis (Walker)<br />
Lep.: Pyralidae 15<br />
Pistiacola cretatus<br />
(Champion) Col.:<br />
Curculionidae 200,205<br />
Pistiacola fasciatus Wibmer<br />
and O'Brien Col.:<br />
Curculionidae 200<br />
Pistiacola sp. nr nigrirostris<br />
Col.: Curculionidae 200<br />
Planococcus citri (Risso)<br />
Hem.: Pseudococcidae 200<br />
Platinglisia noacki Cockerel1<br />
Hem.: Coccidae 101<br />
Platynota rostrana (Walker)<br />
Lep.: Tortricidae 162<br />
platyptera, Calycomyza<br />
pleuralis, Olcella<br />
Plusia oo , see Pseudoplusia<br />
includens 14<br />
Plutella xylostella (Curtis)<br />
Lep.: Yponomeutidae 23<br />
Pochazia antica (Gray)<br />
Hem.: Ricaniidae 121<br />
Podogaster Hym.:<br />
Ichneumonidae 206<br />
Poekilocerus hieroglyphicus<br />
(Klug) Ort.: Acrididae 99<br />
Polychrosis ?carduana, see<br />
hbesia ?carduana<br />
polycoccus, Gibbobruchus<br />
polyphyta, Melangromyza<br />
polyxena, Eryphanis<br />
portulacae, Asphondylia<br />
portulacae, Baris<br />
portulacae, Ceutorhynchus<br />
portulacae, Neolasioptera<br />
portulaecae, Lasioptera<br />
Procecidochares Dip.:<br />
Tephritidae 39<br />
Procecidochares connexa<br />
Macquart Dip.: Tephritidae<br />
40,44,51<br />
procula, Eueides<br />
Prodenia eridania, see<br />
Spodoptera eridania<br />
prolixa, Teleonemia<br />
Promecops Col.:<br />
Curculionidae 29,30, 144<br />
Promecops campanulicollis<br />
Voss Col.: Curculionidae<br />
162<br />
Prosapia bicincta fraterna<br />
(Uhler) Hem.: Cercopidae<br />
89<br />
proserpina, Tarophagus<br />
prosternalis, Diplacaspis<br />
Protonectarina sylveiriae<br />
(Saussure) Hym.: Vespidae<br />
141<br />
Proxenus hennia Swinhoe<br />
Lep.: Noctuidae 202<br />
proxirna, Agromyza<br />
Psalis pennatula (Fabricius)<br />
Lep.: Lymantriidae 225<br />
Pseudaletia unipuncta, see<br />
Mythimna unipuncta<br />
Pseudoderelomus<br />
baridiiformis Champion<br />
Col.: Curculionidae 127,<br />
131<br />
pseudoinsulata, Pareuchaetes<br />
pseudoinsulata, Pareuchaetes<br />
Pseudonapomyza<br />
philippinensis Spencer<br />
Dip.: Agromyzidae 225<br />
Pseudonapomyza spicata<br />
Malloch Dip.:<br />
Agromyzidae 90<br />
Pseudoplusia includens<br />
(Walker) Lep.: Noctuidae<br />
1 "<br />
14<br />
Psigida walkeri (Grote) Lep.:<br />
Cercophanidae 137, 139,<br />
141,144<br />
Psyllaephagus yaseeni Noyes<br />
Hym.: Encyrtidae 141, 142<br />
Psylopigida walkeri, see<br />
Psigida walkeri<br />
Pteronemobius Ort.:<br />
Gryllidae 224<br />
Ptychamalia perlata<br />
(Warren) Lep.:<br />
Geometridae 162<br />
pulchellus, Neohydronomus<br />
punctifrons, Gesonula<br />
puniceus, Acanthoscelides<br />
pusanus, Sogatodes<br />
pusanus, Tagosodes
288<br />
Biological Control of Weeds: Southeast Asian Prospects<br />
Pycnoderes incurvus<br />
(Distant) Hem.: Miridae<br />
128<br />
pygmaeus, Physimerus<br />
quadridentatus,<br />
Acanthoscelides<br />
quadristillata, Echoma<br />
querci, Anoecia<br />
querci, Stegophylla<br />
quinqueguttata, Heliodines<br />
Rastrococcus Hem.:<br />
Pseudococcidae 121<br />
Recilia dorsalis<br />
(Motschulsky) Hem.:<br />
Cicadellidae 89<br />
, recurvalis, Hymenia<br />
recurvalis, Spodoptera<br />
recurvalis, Spoladea<br />
Recurvaria Lep.: Gelechiidae<br />
128<br />
responsalis, Elophila<br />
responsalis, Nymphula<br />
reticulata, Peraphondylia<br />
Rhodobaenus cariniventris<br />
Champ. Col.:<br />
Curculionidae 30,39,47,<br />
51,52<br />
Rhodobaenus<br />
tredecimpunctatus (Illiger)<br />
Col.: Curculionidae 30<br />
Rhopalosiphum nymphaeae<br />
(Linnaeus) Hem.:<br />
Aphididae 77,200,201<br />
Riptortus linearis (Linnaeus)<br />
Hem.: Coreidae 133<br />
ritsemae, Hypolixus<br />
robustae, Liriomyza<br />
rostrana, Platynota<br />
rugosum, Diacamma<br />
russelli, Brachymeria<br />
s-littera, Systena<br />
saccharina, Eldana<br />
Sagra femorata (Drury) Col.:<br />
Chrysomelidae 152<br />
Samea multiplicalis (Guente)<br />
Lep.: Pyralidae 197, 199,<br />
201-203,205<br />
Sameodes Lep.: Pyralidae 76<br />
Sameodes albiguttalis<br />
(Warren) Lep.: Pyralidae<br />
69,73,75-82<br />
Scamurius Hem.: Coreidae<br />
137,139,140,142,143,<br />
145<br />
Sceloenopla Col.:<br />
Chrysomelidae 128, 132<br />
Schizaphis graminum<br />
(Rondani) Hem.:<br />
Aphididae 89<br />
Schizocerella pilicornis<br />
(Holmgren) Hym.:<br />
Tenthredinidae 21 2,213,<br />
217,218<br />
Scotinophara latiuscula<br />
(Breddin) Hem.:<br />
Pentatomidae 57,64, 1 11<br />
scrupulosa, Teleonemia<br />
segnis, Chalcodermus<br />
Selca brunella (Hampson)<br />
Lep.: Noctuidae 120- 123<br />
semicyaneus, Lexyphanes<br />
seminicola, Sibinia<br />
seminis, Geromyia<br />
seminis, Itonida<br />
septentrionalis. Platysimus<br />
serrifer, Homoeocerus<br />
serripes, Chalcodermus<br />
Sesamia Lep.: Noctuidae 225<br />
Sesamia botanephaga Tams<br />
and Bowden Lep.:<br />
Noctuidae 65<br />
Sesamia calamistis Hampson<br />
Lep.: Noctuidae 65<br />
Sesamia cretica Lederer Lep.:<br />
Noctuidae 176<br />
Sesamia penniseti Tams and<br />
Bowden Lep.: Noctuidae<br />
65<br />
setariae, Hysteroneura<br />
sexnebulosa, Leptocysta<br />
seychellarum, Icerya<br />
Sibinia Col.: Curculionidae<br />
144,153<br />
Sibinia aspersa Champion<br />
Col.: Curculionidae 144<br />
Sibinia fatigiata Clark Col.:<br />
Curculionidae 15 1, 155<br />
Sibinia ochreosa Casey Col.:<br />
Curculionidae 15 1, 1 55<br />
Sibinia peruana Pierce Col.:<br />
Curculionidae 15 1, 155<br />
Sibinia seminicola Clark<br />
Col.: Curculionidae 151,<br />
155<br />
Sibinia subulirostris<br />
Hustache Col.:<br />
Curculionidae 144<br />
signoreti, Horciacinus<br />
similis, Chnootriba<br />
similis, Epilachna<br />
similis, Notiphila<br />
similis, Tegosa<br />
simillimus, Odontomachus<br />
simplex, Nisaga<br />
sinuata, Entylia<br />
Sipylus Hem.: Membracidae<br />
121<br />
Sipylus dilatatum (Walker)<br />
Hem.: Membracidae 1 2 1<br />
Sitobion avenue (Fabricius)<br />
Hem.: Aphididae 89<br />
Sitobion leelamaniae (David)<br />
Hem.: Aphididae 94<br />
Sitobion miscanthi<br />
(Takahashi) Hem.:<br />
Aphididae 89<br />
smaragdina, Oecophylla<br />
soccata, Atherigona<br />
socia, Corythuca<br />
Sogatellafurcifera (Horviith)<br />
Hem.: Delphacidae 64, 89<br />
Sogatella kolophon<br />
(Kirkaldy) Hem.:<br />
Delphacidae 176<br />
Sogatella longifurcifera, see<br />
Sogatella vibix<br />
Sogatella panicicola, see<br />
Sogatella vibix<br />
Sogatella vibix (Haupt) Hem.:<br />
Delphacidae 64<br />
Sogatodes pusanus, see<br />
Tagosodes pusanus<br />
solani, Aulacorthum<br />
Solenopsis geminata<br />
(Fabricius) Hym.:<br />
Forrnicidae 1 8 1<br />
solidaginis, Macrosiphum<br />
solidaginis, Uroleucon<br />
solomonensis, Leptocorisa<br />
sorghicola, Contarinia
sororcula, Dioxyna<br />
Sphaeniscus atilus (Walker)<br />
Dip.: Tephritidae 134<br />
Sphenarches anisodactylus<br />
(Walker) Lep.:<br />
Pterophoridae 188, 189<br />
spicata, Pseudonapomyza<br />
Spilosoma virginica<br />
(Fabricius) Lep.: Arctiidae<br />
200,201<br />
spinulosa, Heteropsylla<br />
spiraecola, Aphis<br />
splendida, Melanagromyza<br />
Spoabptera eridania (Stoll)<br />
Lep.: Noctuidae 22, 101,<br />
212<br />
Spoabptera exigua (Hiibner)<br />
Lep.: Noctuidae 22,212<br />
Spodopterafrugiperda (J.E.<br />
Smith) Lep.: Noctuidae 14,<br />
90,225<br />
Spodoptera Iitura (Fabricius)<br />
Lep.: Noctuidae 22,78,<br />
162,167<br />
Spoabptera mauritia<br />
(Boisduval) Lep.:<br />
Noctuidae 78,202<br />
Spoabptera pectinicornis<br />
(Hampson) Lep.:<br />
Noctuidae 197,200,<br />
202-204,206<br />
Spoabptera sunia, see<br />
Neogalea sunia<br />
Spoladea recurvalis<br />
(Fabricius) Lep.:<br />
Pyralidae 23<br />
spretissimum,<br />
Coelocephalapion<br />
squamosus,<br />
Argentinorhynchus<br />
stagnalis, Parapoynx<br />
Staphylus mazans (Reakirt)<br />
Lep.: Hesperiidae 162<br />
Stenchaetothrips biformis<br />
Bagnall Thy.: Thripidae 64<br />
Stegophylla querci (Fitch)<br />
Hem.: Aphididae 89<br />
Stenodiplosis Dip.:<br />
Cecidomyiidae 90<br />
Stenodiplosis panici, see<br />
Contarinia panici<br />
6 Index of scientific names of insects 289<br />
Sternocoelus Col.:<br />
Curculionidae 101<br />
Sternocoelus notaticeps<br />
(Marshall) Col.:<br />
Curculionidae 101<br />
Stiretrus erythrocephalus<br />
Lepeletier and Serville<br />
Hem.: Pentatomidae 29,30<br />
Stomatomyia Dip.:<br />
Tachinidae 1 8 1<br />
strachani, Pinnaspis<br />
striatellus, Laodelphax<br />
strigata, Liriomyza<br />
subornata, Argyractis<br />
subulirostris. Sibinia<br />
sunia, Neogalea<br />
sunia, Spodoptera<br />
Sycanus Hem.: Reduviidae 43<br />
sylveiriae, Protonectarina<br />
Synclita obliteralis (Walker)<br />
Lep.: Pyralidae 201<br />
Syphrea bibiana Bechynk<br />
Jacoby Col.:<br />
Chrysomelidae 15 1<br />
Systena s-littera (Linnaeus)<br />
Col.: Chrysomelidae 144<br />
tabaci, Bemisia<br />
tabaci, Thrips<br />
Tagosodes pusanus (Distant)<br />
Hem.: Delphacidae 64<br />
Tamarixia Eeucaenae Boucek<br />
Hym.: Eulophidae 142<br />
Taractrocera ina Waterhouse<br />
Lep.: Hesperiidae 182<br />
Tarophagus proserpina<br />
(Kirkaldy) Hem.:<br />
Delphacidae 57, 105, 167<br />
taurus, Leptocentrus<br />
Taylorilygus pallidulus<br />
(Blanchard) Hem.: Miridae<br />
1 43<br />
Tegosa claudina<br />
(Eschscholtz) Lep.:<br />
Nymphalidae 129,132<br />
Tegosa similis, see Tegosa<br />
claudina<br />
tegulella, Onebala<br />
Teleonemia Hem.: Tingidae<br />
125,131,132<br />
Teleonemia harleyi<br />
Froeschner Hem.: Tingidae<br />
131<br />
Teleonemia prolixa St31<br />
Hem.: Tingidae 131<br />
Teleonemia scrupulosa Stil<br />
Hem.: Tingidae 132<br />
Teleonemia sp. or spp. nr<br />
prolixa Hem.: Tingidae<br />
127,131<br />
Temelucha ferruginae<br />
(Davis) Hym.:<br />
Ichneumonidae 206<br />
Temnodachrys aphodoides<br />
Col.: Chrysomelidae 144<br />
tenebralis, Nymphula<br />
tenebralis, Parapoynx<br />
tenella, Eurema<br />
Tetraneura basui Hille Ris<br />
Lambers Hem.: Aphididae<br />
89<br />
Tetraneura hirsuta, see<br />
Tetraneura<br />
nigriabdominalis<br />
Tetraneura nigriabdominalis<br />
(Sasaki) Hem.: Aphididae<br />
89<br />
Tetrastichus Hym.:<br />
Eulophidae 1 15, 132<br />
Tetrigella Hem.: Cicadellidae<br />
121<br />
Tettigoniella, see Tettigella<br />
Thagona tibialis (Walker)<br />
Lep.: Lymantriidae 101<br />
Thaia oryzivora Ghauri<br />
Hem.: Cicadellidae 176<br />
theae, Tropicomyza<br />
Thecla azia, see Tmolus azia<br />
Thecla palegon, see Thereus<br />
palegon<br />
Thereus palegon (Cramer)<br />
Lep.: Lycaemidae 129<br />
thetis, Eucerocoris<br />
Thrips hawaiiensis Morgan<br />
Thy.: Thripidae 134<br />
Thrips tabaci Lindeman Thy.:<br />
Thripidae 14, 134<br />
Thrypticus Dip.:<br />
Dolichopodidae 72<br />
Thyanta perditor (Fabricius)<br />
Hem.: Pentatomidae 30
290 Biological Control of Weeds: Southeast Asian Prospects<br />
Thysanoplusia orichalcea Uroleucon Hem.: Aphididae walkeri, Psigida<br />
(~abicius) Lep.:<br />
Noctuidae 3 1<br />
tibialis, Thagona<br />
Tmolus azia (Hewitson) Lep.:<br />
Lycaenidae 145,162<br />
Tomaspis entreriana Berg<br />
Hem.: Cercopidae 143<br />
Toxoptera graminum, see<br />
Schizaphis graminum<br />
trachypterus, Chortogonus<br />
tredecimpunctatus,<br />
Rhodobaenus<br />
triangula, Cicadulina<br />
triangulifer, Hyalomyodes<br />
Tricentrus Hem.:<br />
Membracidae 1 2 1<br />
Trichogramma Hym.:<br />
30<br />
Uroleucon ambrosiae<br />
(Thomas) Hem.:<br />
Aphididae 224<br />
Uroleucon cornpositae<br />
(Theobald) Hem.:<br />
Aphididae 188<br />
Uroleucon solidaginis<br />
(Fabricius) Hem.:<br />
Aphididae 13<br />
ustulata, Ceresa<br />
Valtissius Hem.: Lygaeidae<br />
200<br />
vanillae, Agraulis<br />
variegatus, - Zonocerus<br />
venegasiae, Liriomyza<br />
Xanthaciura insecta (Loew)<br />
Dip.: Tephritidae 14,3 1,<br />
128<br />
xanthaspis, Exorista<br />
Xyloplothrips Thy.:<br />
Phlaeothripidae 134<br />
xylostella, Plutella<br />
xyonysius Hem.: Lygaeidae<br />
132<br />
Xyonysius basalis (Dallas)<br />
Hem.: Lygaeidae 128<br />
Xyonysius inaequalis, see<br />
Xyonysius basalis<br />
Xyonysius sp. nr ementitus,<br />
see Xyonysius basalis<br />
yaseeni, Psyllaephagus<br />
Trichogrammatidae 1 15<br />
Trichotaphe, see Dichomeris<br />
trifolii, Liriomyza<br />
tripsaci, Dalbu lus<br />
Tropicomyia theae (Cotes)<br />
Dip.: Agromyzidae 188<br />
trunculatus, Hypolixus<br />
turbata, Nymphula<br />
turbata, Parapoynx<br />
turrita, Acrida<br />
undalis, Hellula<br />
unipunctata, Mythimna<br />
unipunctata, Pseudaletia<br />
unipunctatus, Germalus<br />
upalusalis, Pionea<br />
urichi, Liothrips<br />
versicolor, Autoba<br />
versicolor, Eublemma<br />
versurella, Coleophora<br />
Vesiculaphis pieridis Basu<br />
Hem.: Aphididae 13<br />
vibix, Sogatella<br />
vinitor, Nysius<br />
virescens, Heliothis<br />
virescens, Nephotettix<br />
virgata, Ferrisia<br />
virginica, Spilosoma<br />
viridiaeneus, Criptocephalus<br />
viridigrisea, Austroasca<br />
viridipennis, Chalcophana<br />
vitripennis, Hyalopeplus<br />
vulgaris, Dialeurodes<br />
zacconius, Chilo<br />
Zatropis Hym.: Pteromalidae<br />
130<br />
zea, Helicoverpa<br />
zebratus, Acanthoscelides<br />
Zizeeria knysna (Trimen)<br />
Lep.: Lycaenidae 22<br />
Zizeeria krupta (Trimen)<br />
Lep.: Lycaenidae 22<br />
Zonocerus variegatus<br />
(Linnaeus) Ort.: Acrididae<br />
13,37<br />
Zulia entreriana, see<br />
Tomaspis entreriana<br />
Zygina Hem.: Cicadellidae<br />
202
General index<br />
(Bold type indicates the pages alternifolius, Mariscus<br />
for the main weeds of altissima, Ageratina<br />
Chapter 4) Amaranthaceae 3,20,21,23,<br />
abaca (Musa textilis) 1 10<br />
Acacia mearnsii 1 41,<br />
143-1 45<br />
Acacia riparia 132<br />
Acalitus 125,127,129<br />
Acalitus adoratus 41,45-48<br />
Acalypha 100<br />
Acremonium zonatum 75,76,<br />
79,82<br />
adenophora, Ageratum<br />
adoratus, Acalitus<br />
Aecidium tithymali 105<br />
aegyptium, Dactyloctenium<br />
Aeschynomene sensitiva 161,<br />
162<br />
afine, Melastoma<br />
agerati, Cercospora<br />
Ageratina altissima 52<br />
Ageratum adenophora 52<br />
Ageratum conyzoides 1,3,<br />
10-16,31,33,50,52,91<br />
Ageratum riparia 52<br />
Ageratum vein yellowing 16<br />
agor 1 10<br />
Agropyron repens 65<br />
aguifigay 222<br />
alang-alang (Imperata<br />
cylindrica) 37<br />
alba, Basella<br />
albida, Mimosa<br />
albilineans, Xanthornonas<br />
Albugo bliti 23<br />
Albugo portulaceae 21 4<br />
Albugo portulacearum 21 4<br />
album, Chenopodium<br />
albus, Amaranthus<br />
alikbangon 56<br />
allelopathy 37, 46, 174,230<br />
Alternanthera philoxeroides 5<br />
Alternaria 98, 100, 1 16<br />
Alternaria compacta 23<br />
Alternaria eichhorniae 77,78<br />
Alternaria passiforae 189<br />
Alternaria tenuis 189<br />
24,216<br />
amaranthi, Aposphaeria<br />
amaranthi, Cacodera<br />
amaranthi, Cercospora<br />
Amaranthus 20-23<br />
Amaranthus albus 24<br />
Amaranthus cruentus 2 1<br />
Amaranthus dubius 21<br />
Amaranthus<br />
hypochondriachus 2 1<br />
Amaranthus oleosa 22<br />
Amaranthus spinosus l,3,<br />
1&24,30,214<br />
Amaranthus tricolor 22,23<br />
Amaranthus viridis 2 1-23<br />
Amblyseius glorius 72<br />
Amblyseius pederosus 72<br />
americanum, Pennisetum<br />
Amphobotrys ricini 100,105<br />
Anacamseros 2 1 8<br />
Andropogon annulatus 89<br />
Andropogon vulgare 94<br />
Andropogoneae 224<br />
anemone 2 1 4<br />
anemone brown ring 21 4<br />
anemone mosaic 16<br />
Anhellia niger 41<br />
anil, Indigofera<br />
annulatus, Andropogon<br />
annuus, Helianthus<br />
anthelminthicum,<br />
Chenopodium<br />
Anthemidae 33<br />
anthocyanin 70<br />
antidotale, Panicum<br />
ants 43,45-47,50,52<br />
anuus, Helianthus<br />
Aphelenchoides fragariae 15,<br />
170<br />
apoda, Phakospora<br />
apoe apoe 198<br />
aponapon 198<br />
Aposphaeria arnaranthi 23<br />
aquatica, Ipomoea<br />
ara tanah 104<br />
Araceae 3, 198<br />
Arctoteae 33<br />
arenaria thamesis,<br />
Meloidogyne<br />
arenaria, Meloidogyne<br />
Arundo donax 8<br />
arvensis, Digera<br />
arvensis, Mentha<br />
Ascochyta 225<br />
Ascochyta portulaceae 2 1 4<br />
asper, Sonchus<br />
asperatg, Mimosa<br />
Aspidosperma ramiflorum<br />
100<br />
Aster31,33,214<br />
aster yellows 32,214<br />
Asteraceae 3, 12,28-31,33,<br />
36,38,50,52,<br />
126,128-130,132<br />
Astereae 33<br />
asthma plant (Euphorbia<br />
hirta) 104<br />
atramentosa,<br />
Myriogenospora<br />
australis, Emex<br />
Austroeupatorium<br />
inulaefolium 14,30,52<br />
Avena, 227<br />
Azolla 7 1<br />
Azolla caroliniana 205<br />
Azolla pinnata 1 15,206<br />
azurea, Eichhornia<br />
ba bawagan 1 10<br />
baccata, Wulfla<br />
baccifera, Melocanna<br />
bamboo 8<br />
Bambusa 8,227<br />
banana 57,63,110,160,180,<br />
21 4<br />
banana poka (Pmsifora<br />
tripartita) 187<br />
bandotan 12<br />
banla saet 138<br />
barley (Hordeum) 8,89,92,<br />
93
292 Biological Control of Weeds: Southeast Asian Prospects<br />
barley stripe mosaic 66<br />
barnyard grass (Echinochloa<br />
crus-galli) 62<br />
Basella alba 22<br />
bayam duri 20<br />
bayokibok 62<br />
bean 88,99<br />
beda bin 70<br />
beet curly top 2 14<br />
Begonia 1 0 1<br />
benghalensis, Commelina<br />
bbo chi 198<br />
Beta vulgaris var. cicla 2 1 1,<br />
212<br />
betae f.sp. portulaceae,<br />
Polymyxa<br />
bicolor, Cochliobolus<br />
. bicolor, Helminthosporium<br />
bicolor, Sorghum<br />
Bidens 29,3 1<br />
Bidens mosaic 32<br />
Bidens mottle 16<br />
Bidens pilosa l,3,26-33,49,<br />
52,91,129-131,181<br />
Bidens witches broom 32<br />
bidenticola. Uromyces<br />
bidentis, Cercospora<br />
biga bigaan 166<br />
Bignoniaceae 5 1<br />
Bipolaris indica 24,2 14<br />
Bipolaris nodulosa, see<br />
Helminthosporium<br />
nodulosum<br />
Bipolaris papendor-i 194<br />
Bipolaris perotidis 225<br />
Bipolaris portulaceae,<br />
see Helminthosporium<br />
portulaceae<br />
Bipolaris setariae 86,91<br />
birds 43,46<br />
biserrata, Nephrolepis<br />
bizat 36<br />
black pepper (Piper nigrum)<br />
100,122,214<br />
black wattle (Acacia<br />
mearnsii) 141,143-1 45<br />
blady grass (Imperata<br />
cylindrica) 193<br />
bliti, Albugo<br />
blue panic (Panicum<br />
antidotab) 175<br />
bb xit 12<br />
borhg 138<br />
botrys, Chenopodium<br />
Brachiaria subquadripara 89<br />
brachiata, Cercospora<br />
brAnjiingAn 222<br />
brassicae, Tylenchorhynchus<br />
braziliensis, Euphorbia<br />
Brevipalpus obovatus 15<br />
Brillantaisia laminum 188<br />
broad sword fern<br />
(Nephrolepis biserrata)<br />
170<br />
buah tikus 186<br />
Buddleia variabilis 101<br />
buffalo grass 180<br />
bulak manok 12<br />
bulbosum, Panicum<br />
bulbous panic (Panicum<br />
bulbosum) 175<br />
bullatum, Tolyposporium<br />
Cacodera amaranthi 23<br />
cactus 214<br />
caerulia, Passiflora<br />
Caesalpiniaceae 139<br />
cajan, Cajanus<br />
Cajanus cajan 16 1, 162,188<br />
Cajanus indicus, see Cajanus<br />
cajan<br />
Calacarus 40,41<br />
Caladium 78<br />
Calendula 3 1<br />
Calenduleae 33<br />
Calliandra selloi 144, 145<br />
camara, Lantana<br />
campestiis, Cuscuta<br />
cancriformis, Triops<br />
canescens, Cercospora<br />
capansa, Euphorbia<br />
capillare, Panicum<br />
capsicum 91,2 14<br />
caracasana, Fleischmannia<br />
caracasana, Wedelia<br />
Cardueae 33<br />
Carlineae 33<br />
caroliniana, Azolla<br />
carpesioides, Venegasia<br />
Carthamnus 33<br />
Caryota 100<br />
cashew 37<br />
cassava (Manihot esculenta)<br />
28,57,63,99,100,181<br />
Cassia 100<br />
cassiicola, Corynespora<br />
Cassytha filiformis 32<br />
cebbensis, Kordyana<br />
cenchroides, Pennisetum<br />
Cercospora 41,167,225<br />
Cercospora agerati 16<br />
Cercospora amaranthi 24<br />
Cercospora bidentis 32<br />
Cercospora brachiata 24<br />
Cercospora canescens 145<br />
Cercospora eupatorii 41<br />
Cercospora eupatorii-<br />
ohratii 41<br />
Cercospora eupatoriicola 41<br />
Cercospora fusimaculans 225<br />
Cercospora marsileae 1 16<br />
Cercospora megalopotamica<br />
32<br />
Cercospora piaropi 77,82<br />
Cercospora portulaceae 2 1 4<br />
Cercospora rodmanii 69,73,<br />
75,77,82,83<br />
Cercospora rottboelliae 226<br />
Cercosporella dominicana<br />
214<br />
Cercosporidium helleri 230,<br />
23 1<br />
cereal chlorotic mottle 93<br />
cespitosa, Euphorbia<br />
chaetochloae, Puccinia<br />
chak thorn 198<br />
Chasmopodium 223<br />
Chenopodiaceae 2 16<br />
Chenopodium 2 1,22<br />
Chenopodium album 22,23<br />
Chenopodium<br />
anthelminthicum 22<br />
Chenopodium botrys 22<br />
cheroma 126<br />
chhlong 174<br />
chi yop luang 148<br />
chili 2 14<br />
chili vein banding 2 14<br />
chinensis, Cuscuta<br />
Chloridoideae 86<br />
Chloris gayana 9 1<br />
chok 198<br />
Chromolaena 37.52
Chromolaena inulaefolium 52<br />
Chromolaena iresinoides 14<br />
Chromolaena ivaefolia 14,<br />
30,48-52<br />
Chromolaena jujuensis 47,<br />
49,50<br />
Chromolaena laevigata 52<br />
Chromolaena microstemon<br />
50<br />
Chromolaena odorata l,3,<br />
13, 14,22,24,30,31,<br />
33,3443,128-132<br />
Chrysanthemum 3 1,33<br />
chrysopids 47<br />
chuntul phnom 1 14<br />
Cichorium 33<br />
Cichorium intybus 15<br />
ciliaris, Digitaria<br />
Cinchona 13 1,132<br />
Cineraria 3 1,33<br />
cinereum, Physarum<br />
Cionothrix praelonga 38,41<br />
citrus 22,28,63, 100, 104,<br />
160,180<br />
clandestinum, Pennisetum<br />
Claviceps 175<br />
clover 2 14<br />
clover big vein 21 4<br />
clover fern (Marsilea minuta)<br />
114<br />
cb c6t heo 12<br />
cb hoi 36<br />
cb l6ng viit 62<br />
cb m2n Mh 86<br />
cb 6ng 174<br />
cb sua l6ng 104<br />
cb trinh nu m6c 138<br />
cobbler's pegs (Bidens<br />
pilosa) 28, 1 30<br />
cochinchinesis, Rottboellia<br />
Cochliobolus bicolor 226<br />
Cochliobolus cymbopogonis<br />
226<br />
Cochliobolus heterostrophus<br />
226<br />
Cochliobolus lunatus 65<br />
Cochliobolus nodulosus, see<br />
Helminthosporium<br />
nodulosum<br />
cocoa 37,127,174,18 1<br />
coconut 28,37,127,139,<br />
160,174,180,186<br />
coffee 28,30,3 1,37,63,98,<br />
100,101,160,180<br />
Coix lacryma-jobi 8<br />
Colletotrichium 16, 133,224,<br />
226<br />
Colletotrichium<br />
gloeosporioides f.sp.<br />
clidemiae 188<br />
Colletotrichium<br />
gloesporioides 133,149,<br />
189<br />
Colletotrichium graminicola,<br />
see Glornerella<br />
grminicola<br />
Colocasia 78<br />
Colocasia esculenta 79<br />
colona, Echinochloa<br />
coloratum, Panicum<br />
Commelina 56,57<br />
Commelina benghalensis 3,<br />
54-58,91,111<br />
Commelina difisa 58<br />
Commelina robusta 57<br />
Commelinaceae 3,56<br />
common heliotrope<br />
(Heliotropiurn europaeum)<br />
131<br />
common sensitive plant<br />
(Mimsa pudica) 160<br />
compacts, Alternaria<br />
compressa, Rottboellia<br />
Coniothyrium 226<br />
conjugatum, Paspalum<br />
conoclinii, Puccinia<br />
conyzoides, Ageratum<br />
coolah grass (Panicum<br />
coloratum) 175<br />
coracana, Eleusine<br />
cordata, Mikania<br />
cordata, Pontederia<br />
cordifolia, Mikania<br />
Coreopsidae 29,32,33<br />
Coreopsis 29,33<br />
corn leaf gall 93,227<br />
corn stunt 93<br />
Corticium sasakii 58,91, 1 1 1<br />
Corynespora cassiicola 1 39,<br />
1 42<br />
Cosmos 33<br />
cotton 20,63,99, 104, 160<br />
couch grass (Cynodon<br />
dactylon) 4<br />
7 General index 293<br />
coumarin 12<br />
cowpea 20,142<br />
crassipes, Eichhornia<br />
creeping panic grass<br />
(Panicum repens) 174<br />
creeping sensitive plant<br />
(Mimosa invisa) 137,138<br />
crenata, Marsilea<br />
Criconemella onoensis 1 1 1<br />
Criconemella xenoplax 2 1 4<br />
Crotolaria 10 1<br />
crotolariae, Phaeotrichoconis<br />
Croton lobatus 15,91<br />
crowsfoot grass (Eleusine<br />
indica) 86<br />
cruentus, Amaranthus<br />
crus-galli var. frumentacea,<br />
Echinochloa<br />
crus-galli, Echinochloa<br />
crusgalli, Ustilago<br />
Ctenopharyngodon idella 72,<br />
75,76<br />
cubensis, Diabole<br />
cucullata, Salvinia<br />
cucumber 24,214<br />
cucumber mosaic 24,189,<br />
214<br />
cucumeris, Thanatephorus<br />
Curvularia 133,223,226<br />
Curvularia cymbopogonis,<br />
see Cochliobolus<br />
cymbopogonis<br />
Cuscuta cmpestris 135<br />
Cuscuta chinensis 135<br />
cylindrica, Impera ta<br />
Cylindrocladium<br />
quinqueseptatum 16, 105<br />
Cymbopogon 8<br />
cymbopogonis, Cochliobolus<br />
cymbopogonis, Curvularia<br />
Cynara 33<br />
Cynara scolymus 33<br />
Cynodon227<br />
Cynodon dactylon 3,4,15,<br />
58,64,88-91,111<br />
cyparissias, Euphorbia<br />
Cyperaceae 3, 1 10<br />
Cyperus iria 9 1<br />
Cyperus rotundus 88,224
294<br />
Biological Control of Weeds: Southeast Asian Prospects<br />
Dactyloctenium 86,92,94<br />
Dactyloctenium aegyptium<br />
90,92,224<br />
Dactyloctenium radulans 92,<br />
94<br />
dactylon, Cynodon<br />
Dahlia 33,50<br />
daoen kaget kaget 160<br />
dasheen<br />
(Colocasia esculenta) 79<br />
daua daua 62<br />
Davalliaceae 170<br />
dayflower (Commelina<br />
benghalensis) 56<br />
delvii, Heterodera<br />
den gai 20<br />
Dendrocalamus 8<br />
Dendrographium<br />
lucknowense 2 14<br />
destructor, Ditylenchus<br />
devil weed (Chromolaena<br />
odorata) 36<br />
Diabole cubensis 149, 150,<br />
151, 155, 156<br />
Diaporthe 226<br />
dichotomiflorum, Panicum<br />
Dichotomophthora indica 2 14<br />
Dichotomophthora lutea,<br />
see Dichotomophthora<br />
indica<br />
Dichotomophthora<br />
portulaceae 21 4<br />
dictyoides, Drechslera<br />
difisa, Commelina<br />
dzfsusa, Marsilea<br />
Digera arvensis 2 1,22<br />
Digera mosaic 24<br />
Digitaria 90,227<br />
Digitaria ciliaris 91<br />
Digitaria sanguinalis 63,224<br />
dilatatum, Paspalum<br />
Diplodia 226<br />
dipsaci, Ditylenchus<br />
distichum, Paspalum<br />
Ditylenchus destructor 91<br />
Ditylenchus dipsaci 2 1 4<br />
djaringan ketul 28<br />
djawan 62<br />
Doassansia 167<br />
dominicana, Cercosporella<br />
donax, Arundo<br />
Drechslera dictyoides 65<br />
Drechslera gigantea 91<br />
Drechslera hawaiiensis 58<br />
Drechslera indica, see<br />
Bipolaris indica<br />
Drechslera maydis, see<br />
Cochliobolus<br />
heterostrophus<br />
Drechslera setariae, see<br />
Bipolaris setariae<br />
dubius, Amaranthus<br />
dumetorum, Schizostachyum<br />
duri semalu 138<br />
2c2ng 70<br />
echeng padi 166<br />
echinatum, Pithecoctenium<br />
Echinochloa 63,227<br />
Echinochloa colona 63-65,<br />
89,91,93<br />
Echinochloa crus-galli 3,57,<br />
60-66 89,91,111,166<br />
Echinochloa crus-galli var.<br />
fiumentacea 62<br />
Echinochloa glabrescens 93<br />
Echinochloa oryzicola 63<br />
Echinochloa pic ta 63.65<br />
Echinochloa utilis 65<br />
Echinochloa walteri 65<br />
Echinopsideae 33<br />
Echium plantagineum 13 1<br />
edulis var. flavicarpa,<br />
Passiflora<br />
edulis, Passiflora<br />
eggplant 2 15<br />
Eichhornia azurea 73, 80, 81<br />
Eichhornia crassipes 1,3,5,<br />
68-83,203,205,206<br />
eichhorniae, Alternaria<br />
eichhorniae, Flechtmannia<br />
eichhorniae, Uredo<br />
ekmaniana, Septoria<br />
ekor kucing 192<br />
elephant grass (Pennisetum<br />
purpureum) 194<br />
Eleusine 86-88,92,94,226,<br />
227<br />
Eleusine coracana 85-89,92,<br />
93<br />
Eleusine indica 1,3,5, 15,<br />
58,84-95,111.224<br />
Eleusine mosaic 89,93<br />
Eleusine tristachya 86<br />
eleusines, Phyllachora<br />
eleusinis, Melanopsichium<br />
eleusinis, Ustilago<br />
Elsinoe 100<br />
Emex australis 130<br />
Emex spinosa 130<br />
encephalitis 71<br />
Entyloma guaraniticum 32<br />
Eragrostis 92<br />
erecta, Tagetes<br />
ergot 18 1<br />
erineum 48,129<br />
erosus, Marsilea<br />
Erythrina lithosperma 30<br />
esculenta, Colocasia<br />
esculenta, Manihot<br />
etjeng padi 70, 166<br />
Euchlaena 90<br />
Eugenia 101<br />
Eupatorieae 33,52<br />
eupatorii, Cercospora<br />
eupatorii, Guignardia<br />
eupatorii-formosani,<br />
Pseudocercospora<br />
eupatorii-odoratii,<br />
Cercospora<br />
eupatoriicola, Cercospora<br />
eupatoriicola, Phomopsis<br />
eupatoriicola, Phyllosticta<br />
Eupatorium 13,5 1<br />
Eupatorium hookerianum,<br />
see Chromolaena jujuensis<br />
Eupatorium odoratum,<br />
see Chromolaena odorata<br />
Euphorbia 97,99,101<br />
Euphorbia braziliensis 101<br />
Euphorbia capansa 100<br />
Euphorbia cespitosa 101<br />
Euphorbia cyparissias 99<br />
Euphorbia geniculata, see<br />
Euphorbia heterophylla<br />
Euphorbia heterophylla 3,91,<br />
96-101,104<br />
Euphorbia hirta 3, 15,99,<br />
102-106<br />
Euphorbia hirtella 100<br />
Euphorbia mosaic 100<br />
Euphorbia ovalifolia 101<br />
Euphorbia pilulifera, see<br />
Euphorbia hirta
Euphorbia prunifolia, see<br />
Euphorbia heterophylla<br />
Euphorbia pseudovirgata 99<br />
Euphorbia pulcherrima 99,<br />
101<br />
Euphorbiaceae 3,98, 100,<br />
101,104<br />
euphorbiae, Uromyces<br />
europaeum, Heliotropium<br />
exaltata, Rottboellia<br />
exigua, Meloidogyne<br />
Exserohilum paspali 182<br />
feather Pennisetum<br />
(Pennisetum polystachion)<br />
192<br />
Ficus 77<br />
filariasis 71<br />
fili2iformis, Cassytha<br />
Fimbristylis littoralis, see<br />
Fimbristylis miliacea<br />
Fimbristylis miliacea 3,57,<br />
58,91,108-111<br />
finger millet (Eleusine<br />
coracana) 85-87<br />
Flechtmannia eichhorniae 72<br />
Fleischmannia caracasana<br />
3 1<br />
Fleischmannia microstemon<br />
14<br />
fluitans, Panicum<br />
foetida, Passiflora<br />
fontana, Montia<br />
foxtail millet (Setaria italica)<br />
8<br />
fragariae, Aphelenchoides<br />
French weed 36<br />
fujikuroi, Gibberella<br />
Fusarium 145<br />
Fusarium moniliforme 226<br />
Fusarium oxysporum f.sp.<br />
elaeidis 24, 41<br />
Fusarium oxysporum f.sp.<br />
passiflorae 188,189<br />
Fusarium roseum 65<br />
fusimaculans, Cercospora<br />
Galinsoga parvi'ora 9 1<br />
garden spurge (Euphorbia<br />
hirta) 104<br />
gatas gatas 104<br />
gayana, Chloris<br />
gelang 10<br />
gelang pasir 210<br />
gelang susu 104<br />
gbndong hcok 104<br />
geniculata, Euphorbia<br />
gewor 56<br />
giant sensitive plant (Mimosa<br />
pigra) 148<br />
Gibberellafujikuroi, see<br />
Fusarium moniliforme<br />
gigantea, Drechslera<br />
Gigantochloa 8<br />
glabrescens,. Echinochloa<br />
glauca, Setaria<br />
glaucum, Pennisetum<br />
globe artichoke (Cynara<br />
scolymus) 33<br />
Gloeocercospora 194<br />
gloeosporioides f.sp.<br />
clidemiae, Colletotrichium<br />
gloeosporioides,<br />
Colletotrichium<br />
Glomerella graminicola 226<br />
glorius, Amblyseius<br />
glumae, Pseudomonas<br />
Glycine max 21 4<br />
glycines, Heterodera<br />
goatweed (Ageratum<br />
conyzoides) 12<br />
goenda 230<br />
golasiman 2 10<br />
goosegrass (Eleusine indica)<br />
86<br />
gooseweed (Sphenoclea<br />
zeylanica) 230<br />
graminicola, Colletotrichium<br />
graminicola, Glomerella<br />
graminicola, Meloidogyne<br />
grandiflora, Portulaca<br />
grass carp<br />
(Ctenopharyngodon idella)<br />
72<br />
grass-like fimbristylis<br />
(Fimbristylis miliacea) 1 10<br />
Grevillea robusta 101<br />
grisea, Magnaporthe<br />
groundnut 20,63,91,93, 104<br />
210,214<br />
groundnut rosette 24,32,57,<br />
93,106,214<br />
7 General index 295<br />
guaraniticum, Entyloma<br />
Guignardia eupatorii 41<br />
guinea grass (Panicum<br />
maximum) 175<br />
hagonoy 36<br />
hairy spurge (Euphorbia<br />
hirta) 104<br />
hairy wandering jew<br />
(Commelina benghalensis)<br />
56<br />
hapla, Meloidogyne<br />
Haplosporella passifloridia<br />
189<br />
hastata, Monochoria<br />
hawaiiensis, Drechslera<br />
hay kai mangda 62<br />
Heleniae 33<br />
Heliantheae 33<br />
Helianthus 24,3 1,33,214<br />
Helianthus annuus 33<br />
Helicotylenchus indicus 2 14<br />
Helicotylenchus multicinctus<br />
15,57,214<br />
Heliotropium europaeum 13 1<br />
helleri, Cercosporidium<br />
Helminthosporium 65,91,98,<br />
100<br />
Helminthosporium bicolor,<br />
see Cochliobolus bicolor<br />
Helminthosporium holmii 91<br />
Helminthosporium maydis 91<br />
Helminthosporium<br />
nodulosum 92<br />
Helminthosporium portulacae<br />
214<br />
Helminthosporium rostratum<br />
194<br />
Helminthosporium turicum<br />
65<br />
Hemphyllium 189<br />
Heterocentron<br />
subtriplinenium 12 1<br />
Heterodera delvii 87,94<br />
Heterodera glycines 2 1 5<br />
Heterodera marioni 21 5<br />
heterophylla, Euphorbia<br />
heterostrophus, Cochliobolus<br />
hibiscus yellow vein mosaic<br />
16,106<br />
hin nu nive tsu bauk 20
296<br />
Biological Control of Weeds: Southeast Asian Prospects<br />
Hirschmaniella 1 1 1, 167,230<br />
Hirschmaniella oryzae 65<br />
Hirschmaniella spinicaudata<br />
9 1<br />
hirta, Euphorbia<br />
hirtella, Euphorbia<br />
holmii, Helminthosporium<br />
hookeriana, Mikania<br />
hookerianum, Eupatorium<br />
Hoplolaimus indicus 2 15<br />
hordeoides, Pennisetum<br />
Hordeum 8,227<br />
hulape 180<br />
Hydrozetes subornata 20 1<br />
Hyparrhenia rufa 3 1<br />
Hypericum perforatum<br />
angustifolium 130<br />
hypochondriachus,<br />
Amaranthus<br />
idella, Ctenopharyngodon<br />
Ilex 101<br />
Impatiens 15<br />
Imperata cylindrica 5,8,24,<br />
37,91,186,193<br />
incognita, Meloidogyne<br />
Indian rhododendron<br />
(Melastoma<br />
rnalabathricum) 120<br />
indica, Bipolaris<br />
indica, Dichotomophthora<br />
indica, Drechslera<br />
indica, Eleusine<br />
indicum, Panicum<br />
indicus, Cajan<br />
indicus, Hoplolaimus<br />
indicus, Pseudocephalobus<br />
Indigofera anil143<br />
intybus, Cichorium<br />
inulaefolium,<br />
Austroeupatorium<br />
Inuleae 33<br />
invadens, Nosema<br />
invisa inermis, Mimosa<br />
invisa, Mimosa<br />
Ipomoea aquatica 78<br />
iresinoides, Chromolaena<br />
iria, Cyperus<br />
Ischaemum rugosum 225<br />
italica, Setaria<br />
itch grass (Rottboellia<br />
cochinchinensis) 222<br />
ivaefolia, Chromolaena<br />
jacobaea, Senecio<br />
jampang canggah 180<br />
Japanese millet (Echinochloa<br />
crus-galli var.<br />
frumentacea) 62<br />
javanica, Meloidogyne<br />
jawg 198<br />
job's tears (Coix lacryma-<br />
jobi) 9<br />
jojoba 2 1 1<br />
jujuensis, Chromolaena<br />
jute 63<br />
ka kiad chrach 166<br />
ka thok rok 186<br />
kak phr6k kdam 1 10<br />
kamplauk 70<br />
kbet choun 210<br />
keladi bunting 70<br />
kelayar 166<br />
kembang gajah 148<br />
keruong padi 174<br />
kiambang besar 198<br />
kikuyu grass (Pennisetum<br />
clandestinum) 193<br />
kirinyu 36<br />
kolokong kabayo 12<br />
Kordyana celebensis 58<br />
krokot 2 10<br />
kumpai jepang 36<br />
Lablab purpureus 188<br />
lablab bean (Lablab<br />
purpureus)<br />
lacryma-jobi, Coix<br />
Lactuca 3 1,33<br />
Lactuca sativa 33<br />
Lactuceae 33<br />
laevigata, Chromolaena<br />
Lagenaria siceraria 188<br />
lamaan 62<br />
laminum, Brillantaisia<br />
lanceolata, Pontederia<br />
Lantana 129<br />
Lantana camara 13,13 1,132<br />
latex 98,99<br />
latifolia, Zizania<br />
Laurus 101<br />
lawn 4,86, 104, 161, 174,<br />
181<br />
legumes 56<br />
Lemna 71<br />
Lemna polyrhiza 1 15<br />
Lemna purpusilla 1 15<br />
lemon grass (Cymbopogon) 8<br />
Leptosphaeria 226<br />
Leptosphaeria proteispora<br />
182<br />
lesser fimbristylis<br />
(Fimbristylis miliacea) 1 10<br />
lettuce (Lactuca sativa) 30<br />
Leucaena leucocephala 141,<br />
142,144,145<br />
leucocephala, Leucaena<br />
Lewisia 218<br />
ligularis, Passifora<br />
lithosperma, Erythrina<br />
littoralis, Fimbristylis<br />
lizards 43,45,46<br />
lobata, Neurolaena<br />
lobatus, Croton<br />
lonchocarpa, Passifora<br />
love-in-a-mist (Passifora<br />
foetida) 186<br />
luc binh 70<br />
lucerne dwarf 66<br />
lucknowense,<br />
Dendrographium<br />
lunatus, Cochliobolus<br />
lutea, Dichotomophthora<br />
luya luyahan 174<br />
luzonense, Panicum<br />
Lyonia ovalifolia 13<br />
mac co 160<br />
Macrophomina phaseolina<br />
100<br />
macrospora, Sclerophthora<br />
Madieae 33<br />
Magnaporthe grisea 87, 92,<br />
226<br />
Magnolia pumila 10 1<br />
mai yah raap yak 148<br />
maiyaraap thao 138<br />
mai yarap 160<br />
mai yarap ton 148<br />
maize (Zea mays) 5,8,20,<br />
28,63-65,80,89-93,<br />
104, 110, 111, 160, 174,<br />
181,210,222,222-226,<br />
maize dwarf mosaic 66,93,<br />
227
maize hoja blanca 227<br />
maize mosaic 227<br />
maize nematode<br />
(Pratylenchus zeae) 91<br />
maize rayado fine 227<br />
maize streak 93<br />
maize stripe 227<br />
maize stripe tenuivirus 227<br />
maize white leaf 227<br />
maize yellow mottle 227<br />
makahiya 160<br />
makahiyang lalaki 138<br />
mala malu 160<br />
malabathricum, Melastoma<br />
malaria 7 1<br />
manatee (Trichechus) 72<br />
mango 20<br />
manicata, Passifora<br />
Manihot esculenta 99<br />
marioni, Heterodera<br />
Mariscus alternifolius 24<br />
Marsilea crenata, see<br />
Marsilea minuta<br />
Marsilea difisa,<br />
see Marsilea minuta<br />
Marsilea erosus, see<br />
Marsilea minuta<br />
Marsilea minuta 3,112116,<br />
167<br />
Marsilea perrieriana, see<br />
Marsilea crenata<br />
Marsilea senegalensis, see<br />
Marsilea minuta<br />
Marsileaceae 3,114<br />
marsileae, Cercospora<br />
max, Glycine<br />
maximum, Panicum<br />
maydis, Drechslera<br />
maydis, Helminthosporium<br />
mayo 104<br />
mays, Zea<br />
mearnsii, Acacia<br />
megalopotamica, Cercospora<br />
Melanopsichium eleusinis 87,<br />
92,94<br />
Melastoma 120<br />
Melastoma afJine, see<br />
Melastoma malabathricum<br />
Melastoma malabathricum 1,<br />
3,118-123<br />
Melastoma polyanthum 121<br />
melastoma (Melastoma<br />
malabathricum) 120<br />
Melastomataceae 3, 120, 121<br />
Melocanna baccifera 8<br />
Meloidogyne 15,32,91, 162,<br />
215,224<br />
Meloidogyne arenaria 15,91,<br />
215<br />
Meloidogyne arenaria<br />
thamesis 15<br />
Meloidogyne exigua 100<br />
Meloidogyne graminicola 65,<br />
91,111,167,176,231<br />
Meloidogyne hapla 32,215<br />
Meloidogyne incognita 15,<br />
23,57,91,105,189,215<br />
Meloidogyne javanica 15,57,<br />
91,100,106,215<br />
Meloidogyne oryzae 1 1 1<br />
melon 99<br />
Mentha arvensis 100<br />
Mexican fire plant<br />
(Euphorbia heterophylla)<br />
98<br />
micrantha, Mikania<br />
microsporida 47,50,52,79<br />
microstemon, Chromolaena<br />
microstemon, Fleischmannia<br />
Mikania 128-1 30,132<br />
Mikania cordata 126<br />
Mikania cordifolia 128-1 32<br />
Mikania hookeriana 13 1<br />
Mikania micrantha l,3,30,<br />
33,52,124-135<br />
Mikania scandens 126<br />
Mikania trinitaria 128, 13 1,<br />
132<br />
Mikania vitifolia 128-1 3 1<br />
mile-a-minute weed (Mikania<br />
micrantha) 126<br />
miliacea, Fimbristylis<br />
miliaceum, Panicum<br />
millet 5,20,63,94<br />
millet panic (Panicum<br />
miliaceum) 175<br />
millet, Japanese (Echinochloa<br />
crus-galli var.<br />
frumentacea) 62<br />
Mimosa<br />
Mimosa albida 144<br />
Mimosa asperata 148<br />
7 General index 297<br />
Mimosa invisa 1,3,136-145,<br />
161,162<br />
Mimosa invisa inermis 139,<br />
141<br />
Mimosa pigra I, 3,138, 140,<br />
142-1 44,146-156,161<br />
Mimosa pigra berlandieri,<br />
see Mimosa asperata<br />
Mimosa pudica l,3, 1 38,<br />
140-142,144,145,<br />
158-162<br />
Mimosa pudica hispida 160<br />
Mimosa pudica unijuga 160<br />
Mimosa quadrivalis 144<br />
Mimosa rixosa 141,144<br />
Mimosa scabrella 143,145<br />
Mimosa somnians 141<br />
Mimosa velloziana 141<br />
Mimosaceae 3,138,139,148,<br />
160<br />
mimosae-invisae, Uredo<br />
mimosae-pigrae,<br />
Phloeospora<br />
mimosine, 149<br />
minuta, Marsilea<br />
minutus, Pratylenchus<br />
Mirabilis 13<br />
mission grass (Pennisetum<br />
polystachion) 192<br />
molesta, Salvinia<br />
mollissima, Passifora<br />
m6nhnyin 1 10<br />
moniliforme, Fusarium<br />
Monochoria hastata 78<br />
Monochoria vaginalis 3,78,<br />
93,115,164-167<br />
monochoria (Monochoria<br />
vaginalis) 166<br />
Montia fontana 2 1 8<br />
Montia pe$oliata 2 1 8<br />
multicinctus, Helicotylenchus<br />
Musa 57<br />
Mutisieae 33<br />
mya by it 2 1 0<br />
Mycovellosiella perfoliata 16,<br />
4 1<br />
mye byet 21 0<br />
myet cho 56<br />
myet ihi 62<br />
myet kha 174<br />
myet ya 222
298 Biological Control of Weeds: Southeast Asian Pro<br />
myet ya nge 222<br />
Myriogenospora atramentosa<br />
182<br />
Myrothecium roridum 78<br />
nam nom raatchasee 104<br />
napier grass (Pennisetum<br />
purpureum) 194<br />
needle burr (Amaranthus<br />
spinosus) 20<br />
Nephrolepidaceae 170<br />
Nephrolepis 17 1<br />
Nephrolepis biserrata 3, 15,<br />
168-171<br />
Neptunia 155<br />
Neptunia oleracea 147, 152,<br />
155<br />
Nerium 100<br />
Neurolaena lobata 13 1<br />
niger, Anhellia<br />
nigrum, Solanum<br />
nodulosa, Bipolaris<br />
nodulosum,<br />
Helminthosporium<br />
nodulosus, Cochliobolus<br />
Nosema 50,206<br />
Nosema nr invadens 8 1<br />
Nosema pilicornis 21 8<br />
notatum, Paspalum<br />
novemnerve, Panicum<br />
novocaledonicus,<br />
Tetranychus<br />
oat pseudorosette 66<br />
oats (Avena sativae) 89,92,<br />
93<br />
obovatus, Brevipalpus,<br />
odorata, Chromolaena<br />
odoratum, Eupatorium<br />
oficinarum, Saccharum<br />
Oidium 149, 162<br />
oil palm 24,28,37, 127, 160,<br />
170,180,193<br />
oleosa, Amaranthus<br />
oleracea, Neptunia<br />
oleracea, Portulaca<br />
oleracea, Spinacia<br />
onoensis, Criconemella<br />
ophiuri, Spacelotheca<br />
ophiuri, Sporisorium<br />
orai 20<br />
Orthogalumna terebrantis 69,<br />
73,75-77,79,80-82<br />
Oryza 227<br />
Oryza sativa 8<br />
oryzae var. commelinae,<br />
Pyricularia<br />
oryzae, Hirschmaniella<br />
oryzae, Meloidogyne<br />
oryzae, Pyricularia<br />
oryzicola, Echinochloa<br />
oryzicola, Eleusine<br />
ovalifolia, Euphorbia<br />
ovalifolia, Lyonia<br />
ovalifolia, Pieris<br />
overeemi, Sporisorium<br />
oxysporum f.sp. elaeidis,<br />
Fusarium<br />
oxysporum f.sp. passiflorae,<br />
Fusarium<br />
paang itik 1 14<br />
painted spurge (Euphorbia<br />
heterophylla) 98<br />
paitan 180<br />
pak prab 56<br />
pak vaen 1 14<br />
paklab 160<br />
pakpawd 230<br />
paku larat 170<br />
palmivora, Phytophthora<br />
paludosum, Panicum<br />
panici,<br />
Parasteneotarsonemus<br />
paniculatum, Talinum<br />
Panicum 91,175,177,227<br />
Panicum antidotale 175<br />
Panicum bulbosum 175<br />
Panicum capillare 175<br />
Panicum coloratum 175<br />
Panicum dichotomiflorum 65<br />
Panicum fluitans 177<br />
Panicum indicum 177<br />
Panicum luzonense 175<br />
Panicum maximum 3 1, 175<br />
Panicum miliaceum 64,90,<br />
175<br />
Panicum novemnerve 175<br />
Panicum paludosum 175<br />
Panicum repens 3,172-177,<br />
181<br />
panicum mosaic 66<br />
papaya 28,142,160,180<br />
papendofli, Bipolaris<br />
Parasteneotarsonemus panici<br />
175,176<br />
Paratylenchus 65<br />
parviflora, Galinsoga<br />
paspali, Exserohilum<br />
paspali, Sorosporium<br />
Paspalum 90, 18 1,227<br />
Paspalum conjugatum 3,89,<br />
178-183<br />
Paspalum dilatatum 18 1<br />
Paspalum distichum 64,93,<br />
181<br />
Paspalum notatum 3 1,89<br />
Paspalum plicatulum 18 1<br />
Pasualum scrobiculatum 18 1<br />
~as>alum vaginatum 1 8 1,<br />
225<br />
paspalum (Paspalum<br />
dilatatum) 18 1<br />
Passiflora 101, 186-189<br />
Passiflora edulis 187<br />
Passiflora edulis var.<br />
flavicarpa 187, 188<br />
Passiflora foetida 3,184-189<br />
Passiflora ligularis 187, 188<br />
Passiflora lonchocarpa 187<br />
Passiflora manicata 187<br />
Passiflora mollissima, see<br />
Passijlora tripartita<br />
Passiflora suberosa 187,188<br />
Passiflora tripartita 185, 187,<br />
188<br />
Passifloraceae 3, 186-1 88<br />
passijlorae, Alternaria<br />
passifloridia, Haplosporella<br />
passion vine butterfly<br />
(Agraulis vanillae) 1 87<br />
passionfruit chlorotic spot<br />
189<br />
passionfruit mosaic 189<br />
passionfruit ringspot<br />
potyvirus 189<br />
passionfruit woodiness<br />
potyvirus 189<br />
pasture 4, 12,28,56,86, 139<br />
181,186<br />
Paterson's curse (Echium<br />
plantagineum) 13 1<br />
pawpaw, see papaya
peanut, see ground nut<br />
pearl millet (Pennisetum<br />
glaucum) 94,193,226<br />
pederosus, Amblyseius<br />
Pellicularia rolfsii 92<br />
penniseti, Spacelotheca<br />
Pennisetum 24, 193, 194,<br />
2 14,227<br />
Pennisetum americanum,<br />
see Pennisetum glaucum<br />
Pennisetum cenchroides 194<br />
Pennisetum clandestinum 193<br />
Pennisetum glaucum 94, 193<br />
Pennisetum hordeoides 1 92<br />
Penniseam polystachion 3,<br />
190-194<br />
Pennisetum polystachion<br />
atrichum 192<br />
Pennisetum polystachion<br />
polystachion 192<br />
Pennisetum polystachion<br />
setosum 192<br />
Pennisetum polystachyon, see<br />
Pennisetum polystachion<br />
Pennisetum purpureum 193<br />
Pennisetum setosum, see<br />
Pennisetum polystachion<br />
Pennisetum typhoideum, see<br />
Pennisetum glaucum<br />
pepperwort (Marsilea<br />
minuta) 1 14<br />
peregrina, Veronica<br />
perfoliata, Montia<br />
perfoliata, Mycovellosiella<br />
perforatum angustifolium,<br />
Hypericum<br />
perotidis, Bipolaris<br />
perrieriana, Marsilea<br />
Pestalopsis 149<br />
Pestalotia 133<br />
petunia 16<br />
Phaeoseptoria 226<br />
phak bia yai 2 10<br />
phak kbiat 166<br />
phak khom nam 20<br />
phak pot 230<br />
phak, top chawaa 70<br />
phak waen 1 14<br />
Phakospora apoda 194<br />
phaseolina, Macrophomina<br />
Phaseolus vulgaris 15<br />
philoxeroides, Alternanthera<br />
Phloeospora mimosae-pigrae<br />
147,150,151,155<br />
Phoenix 100<br />
Phoma 41,214<br />
Phoma tropica 24<br />
Phomopsis 1 49<br />
Phomopsis eupatoriicola 41<br />
Phragmites 8<br />
phti banla 20<br />
Phyllachora eleusines 92<br />
Phyllachora sacchari 226<br />
Phyllostachys 8<br />
Phyllosticta eupatoriicola 41<br />
Phyllosticta stratiotes 157<br />
Physarum cinereum 182<br />
Phytomonas 105<br />
Phytophthora palmivora 100,<br />
105,122,214<br />
piaropi, Cercospora<br />
pickerel weed (Pontederia<br />
cordata) 80<br />
picta, Echinochloa<br />
picta, Eleusine<br />
Pieris ovalifolia 13<br />
pigeon pea (Cajanus cajan)<br />
188<br />
pigra, Mimosa<br />
pigweed (Portulaca oleracea)<br />
210<br />
pilicornis. Nosema<br />
pilosa, Bidens<br />
pilosa, Portulaca<br />
pilulifera, Euphorbia<br />
pineapple 20, 104, 139, 160,<br />
170,180<br />
pineapple yellow spot 14, 16<br />
pinnata, Azolla<br />
pis koetjing 138<br />
pisau pisau 28<br />
Pistia stratiotes 1,3,5,7 1,<br />
1 15. 167,196-207<br />
Pistia virus 1 16, 167<br />
Pithecoctenium 5 1<br />
Pithecoctenium echinatum 5 1<br />
plantagineum, Echium<br />
plantain 1 11<br />
plantarii, Pseudomonas<br />
plicatuhm, Paspalum<br />
Poaceae 3,62,86,174, 180,<br />
192,222,225<br />
7 General index 299<br />
poinsettia (Euphorbia<br />
pulcherrima)<br />
pokok tjerman 36<br />
polyanthes, Vernonia<br />
polyanthum, Melastoma<br />
Polygonum 13<br />
polyhedrosis 5 1,187<br />
Polymyxa betae f.sp.<br />
portulaceae 21 4<br />
polyrhiza, Lemna<br />
polystachion atrichum,<br />
Pennisetum<br />
polystachion setosum,<br />
Pennisetum<br />
polystachion, Pennisetum<br />
Pontederia cordata 73,79,80<br />
Pontederia lanceolata 8 1<br />
Pontederia rotundifolia 80<br />
Pontederiaceae 3,70,79,166<br />
Portulaca 24, 209,218<br />
Portulaca grandijlora 21 4,<br />
216,218<br />
Portulaca oleracea 1,3, 1 5,<br />
91,208-218<br />
Portulaca pilosa 21 6<br />
Portulaca quadrifida 2 16<br />
portulaceae, Albugo<br />
portulaceae, Ascochyta<br />
portulaceae, Bipolaris<br />
portulaceae, Cercospora<br />
portulaceae,<br />
Dichotomophthora<br />
portulaceae,<br />
Helminthosporium<br />
Portulacaecae 3,210,216<br />
portulacearum, Albugo<br />
potato 15,63, 1 1 1, 167<br />
potato virus Y 16<br />
praelonga, Cionothrix<br />
pratensis, Pratylenchus<br />
Pratylenchus 57,215<br />
Pratylenchus minutus ,32,<br />
215<br />
Pratylenchus pratensis 1 5,9 1<br />
Pratylenchus zeae 23,9 1<br />
proso (Panicum novemnerve)<br />
175<br />
proteispora, Leptosphaeria<br />
prunifolia, Euphorbia<br />
Pseudocephalobus indicus 23
300<br />
Biological Control of Weeds: Southeast Asian Prospects<br />
Pseudocercospora<br />
eupatorii-formosani 39,41<br />
Pseudomonas glumae 92<br />
Pseudomonas plantarii 92<br />
Pseudomonas solanacearum<br />
16<br />
pseudovirgata, Euphorbia<br />
Puccinia 24, 100, 1 62<br />
Puccinia conoclinii 16<br />
Puccinia chaetochloae 194<br />
Puccinia rottboelliae<br />
223,226<br />
Puccinia substriata 194<br />
pudica hispida, Mimosa<br />
pudica unijuga, Mimosa<br />
pudica, Mimosa<br />
pulcherrima, Euphorbia<br />
pumila, Magnolia<br />
pungens, Zoysia<br />
purpurea, Tephrosia<br />
purpureum, Pennisetum<br />
purpureus, Lablab<br />
purpusilla, Lemna<br />
purslane (Portulaca oleracea)<br />
210<br />
putri malu 1 48<br />
Pyrenochaeta 227<br />
Pyricularia 58, 176<br />
Pyricularia grisea, see<br />
Magnaporthe grisea<br />
Pyricularia oryzae 65,92,<br />
176<br />
Pyricularia owe var.<br />
commelinae 58<br />
quadr$da, Portulaca<br />
quadripara, Brachiaria<br />
quadrivalis, Mimosa<br />
quinqueseptatum,<br />
Cylindrocladium<br />
Radopholus similis 57, 106,<br />
215<br />
radulans, Dactyloctenium<br />
ragi (Eleusine coracana) 86<br />
ragwort (Senecio jacobaea)<br />
131<br />
ramiflorum, Aspidosperma<br />
Ranunculus sceleratus 16<br />
rats<br />
rau mhc 118 thon 166<br />
mu Sam 2 1 0<br />
red gram 22<br />
r&mb&t& 138<br />
reniformis, Rotylenchulus<br />
repens, Agropyron<br />
repens, Panicum<br />
Rhizoctonia 65, 133<br />
Rhizoctonia solani 78, 100,<br />
167<br />
rice (Oryza sativa) 5,8, 12,<br />
20,23,28,56-58,61,<br />
63-65,71,77,80 88-93,<br />
98 101,104,110,<br />
11 1,114,115,149,160<br />
166,167,174,180 , 186,<br />
193, 199,202,210,222,<br />
225 ,226,230<br />
rice dwarf 66<br />
rice leaf gall 93,227<br />
rice orange leaf 93<br />
rice ragged stunt 93, 167<br />
rice root rot nematode 91<br />
rice tungro 66<br />
rice tungro bacilliform 93<br />
rice tungro spherical 93<br />
rice yellow mottle 93<br />
ricini, Amphobotrys<br />
riparia, Acacia<br />
riparia, Ageratum<br />
rixosa, Mimosa<br />
robusta, Commelina<br />
robusta. Grevillea<br />
rodmanii, Cercospora<br />
rolfsii, Pellicularia<br />
rolfsii, Sclerotium<br />
roridum, Myrothecium<br />
roseum, Fusarium<br />
rostratum, Helminthosporium<br />
Rottboellia 223,227<br />
Rottboellia cochinchinensis 3,<br />
92,220-227<br />
Rottboellia compressa 224,<br />
225<br />
Rottboellia exaltata, see<br />
Rottboellia<br />
cochinchinensis<br />
rottboelliae, Cercospora<br />
rottboelliae, Puccinia<br />
rottboelliae, Spacelotheca<br />
rotundifolia, Pontederia<br />
rotundifolia, Salvinia<br />
rotundus, Cyperus<br />
Rotylenchulus reniformis 15,<br />
23,32,91, 100, 106, 11 1,<br />
182,215<br />
rubber 28,37,120,127,139<br />
160, 170, 174, 180, 193,<br />
194<br />
Rudbeckia 3 1<br />
rufa, Hyparrhenia<br />
rugosum, Ischaemum<br />
rumput belulhng 86<br />
rumput berus 192<br />
rumput bukit 1 10<br />
rumput canggah 180<br />
rumput gajah 192<br />
rumput go1 kar 36<br />
rumput jae jae 174<br />
rumput jurig 192<br />
rumput kerbau 1 80<br />
rumput kuning 192<br />
rumput pait 180<br />
rumput sambou 86<br />
rumput tahi ayam 12<br />
rumput tahi berbau 1 10<br />
saab sua 36<br />
sabung sabungan 86<br />
sacchari, Phyllachora<br />
Saccharum 224,227<br />
Saccharum oficinarum 8<br />
Saccharum spontaneum 227<br />
saltwater couch (Paspalum<br />
distichurn or P. vaginatum)<br />
181<br />
Salvinia 77, 1 15, 197,202<br />
Salvinia cucullata 167,<br />
202,<br />
Salvinia molesta 5,71,77,<br />
167,199,202,206<br />
Salvinia rotundifolia 205<br />
sambau 62<br />
sampeas 160<br />
sanguinalis, Digitaria<br />
sasakii, Corticium<br />
sativa, Lactuca<br />
sativa, Oryza<br />
sawah 70<br />
scabrella, Mimosa<br />
scandens, Mikania<br />
sceleratus, Ranunculus
Schizostachyum dumetorum 8<br />
scitaminea, Ustilago<br />
Sclerophthora macrospora 92<br />
Sclerotinia sclerotiorum 100<br />
sclerotiorum, Sclerotinia<br />
Sclerotium rolfsii 16,92<br />
scolymus, Cynara<br />
scrobic (Paspalum<br />
scrobiculatum) 18 1<br />
scrobiculatum, Paspalum<br />
Secale 227<br />
segan 210<br />
selloi, Calliandra<br />
semalu gajah 148<br />
semhggi 1 14<br />
sembung rambat 126<br />
senduduk 120<br />
Senecio jacobaea 13 1<br />
Senecioneae 33<br />
senegalensis, Marsilea<br />
sensitiva, Aeschynomene<br />
Septoria 41,47<br />
Septoria ekrnaniana 41<br />
sesame 14<br />
Setaria 227<br />
Setaria glauca 63<br />
Setaria italica 8<br />
setariae, Bipolaris<br />
setariae, Drechslera<br />
setosum, Pennisetum<br />
Siam weed (Chromolaena<br />
odorata) 36,46<br />
siceraria, Lagenaria<br />
silisilihan 230<br />
silverbeet (Beta vulgaris var.<br />
cicla) 21 2<br />
similis, Raabpholus<br />
sin ngo let kya 86<br />
sin ngo myet 86<br />
Skranquia 145<br />
smao 110<br />
smao bek kbol62<br />
smao choeung tukke 86<br />
Solanaceae 2 16<br />
solanacearum, Pseuabmonas<br />
solani, Rhizoctonia<br />
Solanum nigrum 14,24<br />
somnians, Mimosa<br />
Sonchus 130<br />
Sonchus asper 14<br />
Sonchus yellow net 32<br />
Sorghum 5,8,28,63,89,90,<br />
91,93,104,110,160,210,<br />
222,224,226,227<br />
Sorghum bicolor 8<br />
Sorghum verticillijlorum 93<br />
Sorosporium paspali 1 82<br />
sourgrass (Paspalum<br />
conjugatum) 180<br />
soybean (Glycine mar) 30,99<br />
160,222<br />
soybean mosaic 32<br />
Spaceloma 100<br />
Spacelotheca ophiuri, see<br />
Sporisorium ophiuri<br />
Spacelotheca penniseti 194<br />
Spacelotheca rottboelliae 227<br />
Spanish needle (Bidens<br />
pilosa) 28<br />
Sphenoclea zeylanica 3, 167,<br />
228-231<br />
Sphenocleaceae 3,230<br />
spiders 45,46,52<br />
Spinacia oleracea 22<br />
spinicaudata, Hirschmaniella<br />
spinosa, Emex<br />
spinosus, Amaranthus<br />
spiny amaranth (Amaranthus<br />
spinosus) 20<br />
spiny pigweed (Amaranthus<br />
spinosus) 20<br />
Spirodela 204<br />
spontaneum, Saccharum<br />
Sporisorium ophiuri 22 1,<br />
223,227<br />
Sporisorium overeemi 176<br />
St John's wort (Hypericum<br />
perforatum angustifolium)<br />
130<br />
stinking passionflower<br />
(Passijlora foetida) 186<br />
Straits rhododendron<br />
(Melastoma<br />
malabathricum) 120<br />
stratiotes, Phyllosticta<br />
stratiotes, Pistia<br />
strawberry 157<br />
suberosa, Passzjlora<br />
subornata, Hydrozetes<br />
subquadripara, Brachiaria<br />
substriata, Puccinia<br />
subtriplinenium,<br />
Heterocentron<br />
7 General index 301<br />
sugar beet 2 14<br />
sugarcane (Saccharum<br />
oficinarum) 5,8,20,63,<br />
65,89,90,93, 104, 110,<br />
111, 139, 160, 174, 181,<br />
193,210,222,226<br />
sugarcane mosaic 66,93,182<br />
sugarcane streak 93<br />
sunflower (Helianthus anuus)<br />
33<br />
swamp panic (Panicum<br />
paludosum) 175<br />
sweet potato 30,57<br />
Tageteae 33<br />
Tagetes erecta 24<br />
Talinum paniculatum 2 1 8<br />
Talinum triangulare 21 8<br />
tapah itik 1 14<br />
tapioca mosaic 16, 106<br />
taro (Colocasia) 57,63,71,<br />
79,110<br />
tarragon 2 1 4<br />
tawbizat 36<br />
tea 28, 37, 104, 127, 139,<br />
160,174,180<br />
teak 37,127<br />
tenuis, Alternaria<br />
Tephrosia purpurea 37<br />
terebrantis, Orthogalumna<br />
termites 43<br />
Tetranychus 140<br />
Tetranychus novocaledonicus<br />
23<br />
Tetranychus urticae 15,100<br />
Thanatephorus cucumeris<br />
167<br />
Tiboochina urvilleana 121<br />
timun padang 186<br />
tithymali, Aecidium<br />
toads 45<br />
tobacco 28,30,63,91,214<br />
tobacco broad ring spot 21 4<br />
tobacco bunchy top 24<br />
tobacco etch 21 4<br />
tobacco leaf curl 16, 106<br />
tobacco mosaic 24,2 14<br />
tobacco streak 2 14<br />
Tolyposporium bullatum 66<br />
tomato91, 100, 111, 142<br />
tomato leaf curl 1 6, 106
302 Biological Control of Weeds: Southeast Asian Prospects<br />
tomato spotted wilt 32<br />
tontrem khet 36<br />
torpedo grass (Panicum<br />
repens) 174<br />
Tradescantia 57<br />
Tragia volubilis 101<br />
translucens, Xanthomonas<br />
triangulare, Talinum<br />
Trichechus 72<br />
tricolor, Amaranthus<br />
tricophora, Ustilago<br />
trinh nu nhon 148<br />
trinitaria, Mikania<br />
Triops cancriformis 1 67<br />
tripartita, Passiflora<br />
tristachya, Eleusine<br />
Triticum 8,227<br />
tropica, Phoma<br />
tuk das khla thom 104<br />
tungro 93<br />
turicum, Helminthosporium<br />
Turneraceae 187<br />
Tylenchorhynchus brassicae<br />
215<br />
typhoideum, Pennisetum<br />
ulam tikus 126<br />
ulasiman 21 0<br />
urd bean yellow mosaic 16,<br />
106<br />
Ureab eichhorniae 82<br />
Ureab mimosae-invisae 145<br />
Uromyces bidenticola 32<br />
Uromyces euphorbiae 100<br />
urticae, Tetranychus<br />
urvilleana, Tiboochina<br />
Ustilago 92<br />
Ustilago crusgalli 66<br />
Ustilago eleusinis, see<br />
Melanopsichium eleusinis<br />
92<br />
Ustilago sciraminea 227<br />
Ustilago tricophora 66<br />
utilis, Echinochloa<br />
utilis, Eleusine<br />
vaginalis, Monochoria<br />
vaginatum, Paspalum<br />
Vanda 15<br />
variabilis, Buddleia<br />
vegetables 8,20,28,56,63,<br />
104,181,193,210<br />
velloziana, Mimosa<br />
Venegasia carpesioides 3 1<br />
Verbena 16<br />
Verbesina 3 1<br />
Vemonieae 33<br />
Veronica peregrina 167<br />
verticilliflorwn, Sorghum<br />
viridis, Amaranthus<br />
vitifolia, Mikania<br />
volubilis, Tragia<br />
vulgare, Andropogon<br />
vulgaris var. cicla, Beta<br />
vulgaris, Phaseolus<br />
walteri, Echinochloa<br />
water clover (Marsilea<br />
minuta) 1 1 3-1 15<br />
water hyacinth (Eichhornia<br />
crassipes) 70<br />
water lettuce (Pistia<br />
stratiotes) 198<br />
Wedelia caracasana 1 4<br />
wheat (Triticum) 58,65,<br />
89-93,111,214<br />
wheat rosette 93<br />
wheat streak mosaic 66<br />
white amur<br />
(Ctenopharyngdon idella)<br />
72<br />
wild passionfruit (Passiflora<br />
foetida) 186<br />
witchgrass (Panicum<br />
capillare) 175<br />
Wulfia baccata 14<br />
xB b6ng 230<br />
Xanthium 15<br />
Xanthomonas albilineans 182<br />
Xanthomonas translucens 66<br />
xenoplar, Criconemella<br />
ya hep 180<br />
ya plong 62<br />
ya sap raeng 12<br />
ya tabsua 12<br />
yaa khaehyon chop k92<br />
yaa prong khaai 222<br />
yaa yaang 98<br />
yah chan ah kat 174<br />
yah chanagard 174<br />
yah koen jam khao 28<br />
yah sua mop 36<br />
yah teenka 86<br />
ye padauk 70<br />
Zea 224,227<br />
Zea mays 8<br />
zeae, Pratylenchus<br />
zeylanica, Sphenoclea<br />
Zinnia 16,3 1,33<br />
Zinnia yellow net 16<br />
zonatum, Acremonium<br />
Zoysia pungens 18 1