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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

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