Eleusine indica - ACIAR
Eleusine indica - ACIAR
Eleusine indica - ACIAR
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84 Biological Control of Weeds: Southeast Asian Prospects<br />
<strong>Eleusine</strong> <strong>indica</strong><br />
(after Holm et a/. 1977)
Map 4.8.1 <strong>Eleusine</strong> <strong>indica</strong><br />
4.8 <strong>Eleusine</strong> <strong>indica</strong> 85<br />
<strong>Eleusine</strong> <strong>indica</strong><br />
<strong>Eleusine</strong> <strong>indica</strong> 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. <strong>indica</strong> 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 <strong>Eleusine</strong> <strong>indica</strong> (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. <strong>indica</strong> 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 <strong>Eleusine</strong>, 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. <strong>indica</strong> (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 <strong>Eleusine</strong> and Dactyloctenium are closely related.<br />
E. <strong>indica</strong> 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 <strong>Eleusine</strong> <strong>indica</strong> 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. <strong>indica</strong> 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 <strong>Eleusine</strong> 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 <strong>Eleusine</strong> and its close relatives might well be con-<br />
sidered for biological control of E. <strong>indica</strong> except in India or other regions where finger<br />
millet is an important cereal.<br />
E. <strong>indica</strong> 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. <strong>indica</strong>, 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. <strong>indica</strong> 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. <strong>indica</strong>. 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. <strong>indica</strong><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. <strong>indica</strong> 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 <strong>Eleusine</strong> 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 <strong>Eleusine</strong> 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. <strong>indica</strong> to establish whether or not adequately specific species occur<br />
there. A two year (or longer) survey of E. <strong>indica</strong> in several regions of Africa would prob-<br />
ably be required, together with observations on whether the organisms found attacking<br />
E. <strong>indica</strong> also attack E. coracana, nearby grasses or other plants. The relevant regions for<br />
study in Africa and Madagascar are <strong>indica</strong>ted 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. <strong>indica</strong> 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. <strong>indica</strong> 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. <strong>indica</strong>, 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. <strong>indica</strong><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 />
<strong>Eleusine</strong>. 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 <strong>Eleusine</strong> <strong>indica</strong>: 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, <strong>Eleusine</strong> 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 <strong>Eleusine</strong> <strong>indica</strong> 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 />
<strong>Eleusine</strong> 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 />
<strong>Eleusine</strong> 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. <strong>indica</strong><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. <strong>indica</strong> 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 <strong>Eleusine</strong> <strong>indica</strong>: 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 <strong>Eleusine</strong> <strong>indica</strong> 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. <strong>indica</strong> 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 <strong>Eleusine</strong> <strong>indica</strong>: 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. <strong>indica</strong>,<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. <strong>indica</strong><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, <strong>Eleusine</strong><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. <strong>indica</strong><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 <strong>Eleusine</strong> 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. <strong>indica</strong><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 <strong>Eleusine</strong> 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 <strong>indica</strong>: viruses.<br />
Virus Country Other hosts<br />
cereal chlorotic mottle<br />
corn leaf gall<br />
corn stunt<br />
<strong>Eleusine</strong> 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. <strong>indica</strong> 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 <strong>Eleusine</strong> <strong>indica</strong> 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 <strong>Eleusine</strong> 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 <strong>Eleusine</strong> and Wapshere 1990c,<br />
Dactylocteniurn: Zundel 1953<br />
tentatively identified<br />
from D. radulans in<br />
Queensland, but not<br />
from E. <strong>indica</strong>
4.8 <strong>Eleusine</strong> <strong>indica</strong><br />
Map 4.8.2 Distribution of <strong>Eleusine</strong> <strong>indica</strong> 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), <strong>indica</strong>ting 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