ARTICLE IN PRESS

Crop Protection 23 (2004) 739755

Review

Biology and management of the sugarcane aphid, Melanaphis

sacchari (Zehntner) (Homoptera: Aphididae), in sorghum: a review
B.U. Singh*, P.G. Padmaja, N. Seetharama
National Research Centre for Sorghum, Rajendranagar, Hyderabad, Andhra Pradesh 500 030, India
Received 1 October 2003; received in revised form 3 December 2003; accepted 13 January 2004

Abstract
The sugarcane aphid, Melanaphis sacchari (Zehntner, 1897) is a key pest on sorghum and sugarcane in many areas of Africa, Asia,
Australia, the Far East, and parts of Central and South America. The status of research of its geographical distribution, host range,
nature of damage, extent of crop losses, and ecobiology in sorghum is summarized and research programs in different countries are
reviewed. Numerous germplasm accessions, A/B- and R-lines, agronomic elite lines, hybrids, and varieties, identied as sources of
resistance providing genetic diversity from different countries are listed. Studies on the components of resistance showed the
predominance of antixenosis for colonization/establishment on IS 1144C, IS 12664C, and TAM 428, and antibiosis was observed on
IS 12609C, IS 12664C, and TAM 428 for least number of days to reproduction, greater mortality, shorter longevity, and production
of no or fewer nymphs. The morpho-physiological traits and biochemical factors associated with resistance have been discussed.
There is a signicant decline in diastase activity but increase in crude ber and carbohydrates in the grain due to infestation by M.
sacchari. It is a vector of three persistent viruses (millet red leaf, sugarcane yellow leaf, and sugarcane mosaic viruses). Among the
control tactics, cultural practices, natural enemies, and chemical control together can prevent the sugarcane aphid from reaching the
economic threshold levels. Current progress has been reviewed and ideas for future research are suggested.
r 2004 Elsevier Ltd. All rights reserved.
Keywords: Biology; Management; Mechanisms of resistance,Melanaphis sacchari; Sorghum; Sugarcane aphid; Varietal resistance

Contents
1.

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

740

2.

Geographical distribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

740

3.

Host range . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

740

4.

Nature of damage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

742

5.

Crop losses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

742

6.

Ecobiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

743

7.

Varietal resistance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

743

8.

Virus vector . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

748

9.

Cultural practices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

748

10. Natural enemies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

748

11. Chemical control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

751

*Corresponding author.
E-mail address: singhbu@rediffmail.com (B.U. Singh).
0261-2194/$ - see front matter r 2004 Elsevier Ltd. All rights reserved.
doi:10.1016/j.cropro.2004.01.004

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12. Future research . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

751

Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

752

References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

752

1. Introduction
The sugarcane aphid, Melanaphis sacchari (Zehntner,
1897) (Homoptera: Aphididae) has been reported
variously as Aphis sacchari (Zehntner) (Zimmerman,
1948) and Longiunguis sacchari (Zehntner) (Eastop,
1965). Roy Chaudhuri and Banerjee (1974) synonymized both these species as M. sacchari (Zehntner).
Eastop and Hille Ris Lambers (1976) reviewed the
literature on the nomenclature of sugarcane aphid, M.
sacchari, and listed that the genera, Geoktapia Mordovilko (1921), Longiunguis Van der Goot (1977), Masraphis Soliman (1938), Nevsikia Mordvilko (1932),
Piraphis Borner (1932), Schizaphideilla Hille Ris Lambers (1939), Yezabura Matsumura (1917) as synonyms
of the genus Melanaphis Van der Goot (1917).

Table 1
Geographical distribution of the sugarcane aphid, Melanaphis sacchari
(Zehntner)
Country

Reference

Angola
Argentina
Australia
Bhutan
Botswana
Brazil
China

Mead (1978)
Delno (1985)
Passlow (1985)
Agarwala (1985)
Flattery (1982)
Mead (1978)
Wang (1961), Mead (1978), and Miao and Sunny
(1987)
Mead (1978)
Mead (1978)
Mead (1978)
Megenasa (1982)
Denmark (1988)
Mead (1978)
Mead (1978)
Young (1970), Jotwani and Young (1972), Young
and Teetes (1977), Shuja-Uddin (1975), David and
Sandhu (1976), Alexander and Madhusudhanrao
(1977), Varma et al. (1978), Bapat (1981), Bhagat
(1981), Agarwala et al. (1983), Mote (1983), Mote
and Kadam (1984), Patil (1992), and Balikai (1997)
Mead (1978)
Edward (1937)
Setokuchi (1973), Mead (1978), Hagio et al. (1985),
and Hagio and Ono (1986)
Le Pelley (1959), Nye (1960), and Starks (1969)
Mead (1978)
Hamid (1983)
Mead (1978)
Rueda and Catling (1978), and Mead (1978)
Brain (1929), Muller and Scholl (1958), Matthee and
Oberholzer (1958), Matthee (1962), van Rensburg
and van Hamburg (1975), Anonymous (1981), van
Rensburg and Malan (1993), and van den Berg
(1999)
Schmutterer (1969), and Mead (1978)
Mead (1978)
Chang (1981a, b), Chang and Fang (1984), and
Wilbrink (1922)
Bohlen (1973)
Young (1970), Banzoger (1976), and Meksongsee
and Chawanapong (1985)
Mead (1978)
Schmutterer (1969), and Mead (1978)
Delno (1985)
Mead (1978), Sanchez and Cermeli (1987), and
Aponte et al. (1988)
Sithole et al. (1987)

Colombia
Ecuador
Egypt
Ethiopia
Florida
Haiti
Hawaii
India

2. Geographical distribution
The geographical distribution of M. sacchari follows
the cultivation of sorghum (Sorghum bicolor (L.)
Moench) and sugarcane (Saccharum officinarum (L.))
worldwide covering Angola, Brazil, China, Colombia,
Ecuador, Egypt, Ethiopia, Haiti, Hawaii, India, Indonesia, Japan, Jamaica, the middle East, Nigeria, Pakistan, Peru, Philippines, Sudan, Thailand, Trinidad,
Tabago, Uganda, and Venezuela (Eastop, 1955, 1965;
Mead, 1978; CIE, 1981) (Table 1). It is considered as an
economically important pest on sorghum in China
(Wang, 1961), Taiwan (Chang, 1981a, b; Pi and Hsieh,
1982a), Japan (Setokuchi, 1973), India (Young, 1970),
South Africa (van Rensburg, 1973a), and Botswana,
while it is common on cultivated sorghum in Zimbabwe
(Flattery, 1982), where its economic importance has not
been determined (Page et al., 1985). In North America,
it occurs on sugarcane in Florida (Summers, 1978;
Mead, 1978; Denmark, 1988), Hawaii (Zimmerman,
1948; Pemberton, 1948), and Louisiana (White et al.,
2001), and its economic status on sugarcane still remains
unclear (White et al., 2001).

3. Host range
Although M. sacchari (Zehntner) (Homoptera: Aphididae) is a minor pest on several crops, its pest status has
increased rapidly since the early 1970s. The genus
Melanaphis has 20 species associated with Gramineae

Indonesia
Jamaica
Japan
Kenya
Nigeria
Pakistan
Peru
Philippines
South
Africa

Sudan
Tabago
Taiwan
Tanzania
Thailand
Trinidad
Uganda
Uruguay
Venezuela
Zimbabwe

(Blackman and Eastop, 1984). The host range of M.

sacchari is largely restricted to the species of the genera:
Saccharum, Sorghum, Oryza, Panicum, and Pennisetum

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Table 2
Host range of the sugarcane aphid, M. sacchari (Zehnt.) reported from different countries
Scientic name

Common name

Country from
which reported

Reference

Cynodon dactylon (L.)

Bermuda grass,
Burmagrass,
Common stargrass,
Devilgrass,
Dhubgrass
Ornamental grass,
Japanese silvergrass
Paddy,
Rice
Barnyard grass,
Jungle rice,
Tufted annual grass
Hamilgrass,
Jacquin Guineagrass
Hairy crabgrass

Taiwan

Wilbrink (1922)

Japan

Setokuchi (1973), and Kawada (1995)

China,
USA (Florida)
USA (Florida),
Taiwan

Miao and Sunny (1987)

Denmark (1988)
Denmark (1988)
Wilbrink (1922)

Botswana,
Zimbabwe
USA (Florida)

van Rensburg (1973a)

van Rensburg (1973a)
Wilbrink (1922)

USA (Florida)
Argentina,
USA (Florida),
USA (Hawaii),
India (Sikkim),
India (Tamil Nadu),
India (Uttar Pradesh),
India (West Bengal),
Jamaica,
USA (Louisiana),
Pakistan,
Philippines,
Taiwan
South Africa,
USA (Florida)

Denmark (1988)
Delno (1985)
Mead (1978), and Denmark (1988)
Pemberton (1948)
Agarwala et al. (1983)
Alexander and Madhusudhanrao (1977)
Shuja-Uddin (1975), and Varma et al. (1978)
Agarwala et al. (1983)
Edward (1937)
White et al. (2001)
Hamid (1983)
Rueda and Catling (1978)
Wilbrink (1922)
van Rensburg (1973a, b)
Wilbrink (1922)

Argentina,
USA (Florida),
India (Karnataka),
India (Kashmir),
India (Maharashtra),
India (Punjab),
Japan,

Delno (1985)
Wilbrink (1922), and Denmark (1988)
Patil (1992), and Balikai (1997)
Bhagat (1981)
Mote (1983), and Mote and Kadam (1984)
David and Sandhu (1976)
Setokuchi (1973), Hagio et al. (1985), and Hagio
and Ono (1986)
van Rensburg and van Hamburg (1975)
van Rensburg and Malan (1983)
Chang (1981a, b), and Chang and Fang (1984)
Banzoger (1976)
Delno (1985)
Sanchez and Cermeli (1987), and Aponte et al. (1988)
Kawada (1995)
van Rensburg (1973a)

Miscanthus sinensis, (L.)

Oryza sativa (L.)
Panicum colonum
(Syn: Echinochloa colonum)
Panicum maximum Jacq.
Paspalum sanguinale Lamarck
(Syn: Digitaria sanguinalis)
Pennisetum sp.
Saccharum officinarum

Setaria italica (L.) Beauv.

Sorghum bicolor (L.) Moench

Sugarcane

Boar millet,
Foxtail millet,
German millet,
Hay millet,
Italian millet,
Nunbank setaria
Sorghum

South Africa,

Sorghum halepense (L.) Pers.

Sorghum verticilliflorum (Steud.)

Stapf.
Zea mays (L.)

Aleppo grass,
Aleppo millet grass,
Cuba grass,
Johnson grass
Wild Sudangrass
Maize

Taiwan,
Thailand,
Uruguay,
Venezuela
Japan,
South Africa

South Africa
Bhutan

(Denmark, 1988) (Table 2). Brain (1929) recorded M.

sacchari as a potential pest of sugarcane in coastal areas
of Natal; and Muller and Scholl (1958), Matthee and

van Rensburg (1973a), and van Rensburg and

van Hamburg (1975)
Agarwala (1985)

Oberholzer (1958), and Matthee (1962) found it as a

regular pest on sorghum in Northern Transvaal in South
Africa. In addition, its host range spreads to Setaria

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italica (L.) Beauv. in the USA (Wilbrink, 1922) and

South Africa (van Rensburg, 1973a), and Zea mays (L.)
in Bhutan (Agarwala, 1985), including species of grasses
such as Cynodon dactylon (L.) in Taiwan (Wilbrink,
1922), Miscanthus chinensis (L.) in Japan (Setokuchi,
1973; Kawada, 1995), Paspalum sanguinale Lamarck in
the USA (Wilbrink, 1922), Sorghum halepense (L.) Pers.
in South Africa (van Rensburg, 1973a) and Japan
(Kawada, 1995), and Sorghum verticilliflorum (Steud.)
Stapf. in South Africa (van Rensburg, 1973a; van
Rensburg and van Hamburg, 1975). A list of the host
plants from which the sugarcane aphid has been
reported is provided in Table 2. The sugarcane aphid
is anholocyclic throughout most of its range in tropical
and subtropical regions, but some sexual oviparae have
been observed on Gramineae in Punjab, India (David
and Sandhu, 1976). In addition, monoecious holocyclic
forms have also been reported on sorghum (David and
Sandhu, 1976) and sugarcane (Yadava, 1966). Both alate
and apterous virginoparous adults showed a stronger
tendency of preference to Sorghum bicolor and S. halepense
than Japanese silvergrass, Miscanthus sinensis (Anders)
(Kawada, 1995), but had a strong preference for M.
sinensis over sugarcane (S. officinarum L.).

4. Nature of damage
Damage to sorghum by the sugarcane aphid depends
on a number of factors including aphid density and
infestation duration. Sorghum is typically infested soon
after plant emergence, but signicant infestations
usually occur during late growth stages, and in dry
periods (van Rensburg, 1973a). Sorghum responses to
M. sacchari injury include purple leaf discoloration of
seedlings followed by chlorosis, necrosis, stunting, delay
in owering, and poor grain ll, including quality and
quantity yield losses. The sugarcane aphid feeds on the
abaxial surface of older sorghum leaves. Leaves below
the infected ones are often covered with sooty molds
which grow on the honeydew produced by the aphid
(Narayana, 1975). Plant stress due to drought may
intensify damage to sorghum by the sugarcane aphid.
The importance of M. sacchari as a pest on sorghum
results from its colonization when plants are 23 weeks
old. Despite early colonization, notable increases in
aphid numbers take place only after panicle exsertion
(van Rensburg, 1973a). Therefore, the sugarcane aphid
colony has a relatively narrow time window within
which to increase population buildup. Physiological and
biological changes taking place during sorghum plant
development can cumulatively affects the exponential
growth rates during early and mid-season, reaching as
many as 30 000 aphids on a single plant (Setokuchi,
1977). However, the aphid densities decline quickly in
23 weeks after peak abundance, and the factors

inuencing decline are alate dispersal induced by aphid

density as well as the poor host condition (van
Rensburg, 1973b).
There is a signicant increase in population of M.
sacchari on sorghum from the boot to the soft dough
stage (4070 days after planting) in the spring, and
heading to harvesting (60100 days after planting) in
autumn (Fang, 1990). Waghmare et al. (1995) observed
population increase and peaks during January, when the
post-rainy sorghum crop was between owering and
milk stage, and declined thereafter till maturity. Plant
growth stage and temperature had signicant effects on
the population buildup of the sugarcane aphid, and
dispersal occurs within 610 days at a temperature
regime of 15.1 C and 31.0 C (Balikai, 2001), 16.0 C and
29.0 C (Mote and Kadam, 1984), 19.5 C and 34.7 C
(Narayana et al., 1982), 22.5 C and 32.5 C, and at 84%
RH (AICSIP, 19792003) and 18.031.0 C (van Rensburg, 1973a, b), but the population died at 35.0 C
(Behura and Bohidar, 1983). In addition to the
temperature, cloudy weather together with increasing
humidity can result in aphids colonies completely
covering the abaxial surface of all leaves of sorghum
plants (Mote, 1983). The highest rate of population
buildup was at 94% and 43% RH and at 11.4 C and
30.0 C temperature in the morning and afternoon,
respectively (Waghmare et al., 1995). Aphid density was
greater under irrigation than in unirrigated conditions,
and its occurrence on sorghum at milk stage did not
affect the grain yields severely, but the fodder quality
deteriorated (Balikai, 2001).

5. Crop losses
The sugarcane aphids remove the plant sap from
xylem tissues of leaves, and in large densities they cause
physiological losses such as wilting/curling of leaves,
and also result in chlorosis. However, the aphid
numbers necessary to cause yield reductions in sorghum
vary based on the plant stage, interval between, and
duration of infestation. The degree of plant moisture
stress under which sorghum is grown as well as the
induction of stress due to aphid infestation also plays a
signicant role in the amount of aphid injury that can be
tolerated.
Sorghum yield losses ranging from minor to severe
have been reported in Botswana (Anonymous, 1974;
Flattery, 1982), Zimbabwe (Page et al., 1985) and India
(Mote and Kadam, 1984; Mote et al., 1985). In South
Africa, grain yield losses reached 60% (Matthee, 1962),
and 4678%, without insecticide control (van Rensburg
and van Hamburg, 1975; van Rensburg, 1979; van den
Berg, 2002). There are few direct and indirect estimates
of the sugarcane aphid damage in sorghum and
sugarcane. In sorghum, the losses varied between

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1226% and 1031% with an overall loss of 16% and

15% for grain yield and fodder yield, respectively
(Balikai, 2001). Sugarcane aphid infested sorghum grain
was signicantly associated with the poor preparation of
beverages (Pi and Hsieh, 1982b), and reduction in
diastatic activity, malt, and abrasive hardness index (van
den Berg et al., 2003) as well as causing grain yield
reduction and poor quality of forage sorghums (Setokuchi, 1979) similar to decrease in crude protein and
increase in neutral detergent ber (NDF) by the yellow
sugarcane aphid, Sipha flava (Fukumoto and Mau,
1989). There was a signicant reduction of 40.2% and
39.1% in grain and fodder yields, respectively, when
infested at 60 DAE, compared to 70 and 80 DAE with
23% and 28% loss in grain yield, and 15% and 17%
reduction in forage yields, respectively (Balikai, 2001).

6. Ecobiology
The earliest detailed study of the biology and life
history of the sugarcane aphid has been on sorghum in
South Africa (van Rensburg, 1969, 1973a, b, 1976) and
to a limited extent on sugarcane in Japan (Setokuchi,
1980, 1988) and sorghum in India (Varma et al., 1978).
The sugarcane aphid forms colonies of lemon-yellow
apterae and alate individuals on the abaxial surface of
basal leaves of a sorghum plant. Some alates have
patterned black markings along the dorsal scleritis
(Eastop, 1955; Roy Chaudhuri and Banerjee, 1974;
Blackman and Eastop, 1984). Reproduction is predominantly asexual with adults being either apterae or
alate viviparous females. Sexual reproduction is also
known to occur on sorghum (David and Sandhu, 1976),
however, the environmental conditions under which
sexual reproduction takes place have not been reported.
It has four nymphal stadia, which are completed in 4.3
12.4 (Chang et al., 1982) or in 5 days (Manthe, 1992).
Adults normally survive for 1016 (Meksongsee and
Chawanapong, 1985), 1437 (Chang et al., 1982), or 28
days (van Rensburg, 1973a), and produce up to 68
nymphs female 1 with an average of 34 (Meksongsee
and Chawanapong, 1985), 4589 (Chang et al., 1982), or
96 at 18.031.0 C (van Rensburg, 1973a). Alates
produce fewer nymphs and have a shorter life expectancy (van Rensburg, 1973a). It develops 5161 generations, averaging 56 generations annually under
screenhouse conditions (Chang et al., 1982). The life
span of each generation is shorter in summer than in
winter (Chang et al., 1982). The number of days for
nymphal development was increased with a decrease in
longevity and fecundity in aphids reared on sorghum at
25.0 C and 16 h photoperiod (Kawada, 1995). There is
also evidence that the performance of the sugarcane
aphid varies in its adaptation when reared on sugarcane
alone, while those reared on sorghum are adapted to

743

both sorghum and sugarcane (Setokuchi, 1988). The

population density is inuenced by variable temperatures and rainfall pattern (Chang et al., 1982).
It overwinters parthenogenetically on ratoon sorghum
and wild alternate hosts such as S. verticilliflorum, S.
halepense (L.), Panicum maximum Jacq., and Setaria
spp. (van Rensburg, 1973a). The dispersal of alates
throughout the year ensures that young cultivated
sorghum is infested soon after germination (van
Rensburg, 1973a). The seasonal history of the sugarcane
aphid has been described in two phases: (i) fast
exponential increase in numbers during the early season
is terminated by heavy dispersal in the mid-season, and
responsible for considerable loss in yield; and (ii) late
summer phase, wherein natural enemies regulate the
aphid populations at sub-economic levels for rest of the
season (van Rensburg and van Hamburg, 1975). Since
highly signicant correlations occur between the population development of wheat aphid and the sugarcane
aphid, the developmental parameters of the former has
been suggested to predict the similar population trend of
M. sacchari on sorghum (Niu, 1987).

7. Varietal resistance
Aphid density and plant damage under natural
infestations have been used to select resistant sorghum
genotypes in the greenhouse and eld conditions
(Setokuchi, 1976; Pi and Hsieh, 1982a; Hagio and
Ono, 1986). Seedling and mature plant evaluations
displayed similar results (Pi and Hsieh, 1982a; Hagio
and Ono, 1986), however, seedlings were preferred for
easy handling and control of infestation levels (Teetes,
1980).
Sources and mechanisms of resistance: Several sources,
levels, and mechanisms of resistance in sorghum to the
sugarcane aphid comprising germplasm accessions,
parental lines (A/B and R lines), agronomic elite lines,
hybrids, varieties, and locals have been reported from
different countries (Tables 36). Resistance to insects
has been characterized into three components as
antixenosis (nonpreference), antibiosis, and tolerance
(Painter, 1951; Horber, 1980; Smith, 1989; Smith et al.,
1994). However, antixenosis and antibiosis resistance to
M. sacchari in sorghum do not differ with plant age
(Teetes, 1980). Both alate and apterous virginoparous
adults showed a stronger tendency of preference to S.
bicolor and S. halepense rather than M. sinensis.
Tolerance to the sugarcane aphid injury in sorghum
increases greatly with slight increase in plant height, and
has inherent advantage over antibiosis and antixenosis
in that it does not impose selection pressure on aphid
populations, and thus may have greater permanence.
Setokuchi (1988) reported that M. sacchari failed to
establish on resistant lines under eld conditions

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Table 3
Germplasm accessions of sorghum resistant to the sugarcane aphid, M. sacchari (Zehnt.) reported from different countries
Germplasm accession

Origin

Classication

Resistance
Level

IS
IS
IS
IS
IS
IS

44
84
718
1063
1117
1133C (SC 202)

Country from
which reported

Reference

India
Japan
India
India
India
Botswana,
Zimbabwe
Botswana,
Zimbabwe
Botswana,
Zimbabwe
Botswana,
Zimbabwe
Botswana,
Zimbabwe
Botswana,
Zimbabwe
India
India
Japan
India
Botswana,
Zimbabwe
India
Botswana,
Zimbabwe
Botswana,
Zimbabwe
Botswana,
Zimbabwe
Botswana,
Zimbabwe
Botswana,
Zimbabwe
Japan
Botswana,
Zimbabwe
Botswana,
Zimbabwe
Botswana,
Zimbabwe
Botswana,

Balikai (1993)
Hagio and Ono (1986)
AICSIP (1988)
AICSIP (1988)
AICSIP (1988)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Mote and Shahane (1988, 1993)
AICSIP (1997)
Hagio and Ono (1986)
Balikai (1993)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
AICSIP (1980)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Hagio (1987, 1992) and Hagio and Ono
(1986)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and
Manthe (1992) and
Manthe (1992) and
Manthe (1992) and
Chang (1981a, b)
Manthe (1992) and
Manthe (1992) and

Manthe (1992) and Teetes et al. (1995)

Hagio and Ono (1986), Hagio et al. (1985),
and Hagio (1987, 1992)
Wenzell et al. (1998)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)

Mechanism

USA
Mexico
USA
India
India
India

Bicolor
Durra/Kaura
Bicolor
Durra
Durra
Durra

R
MR
R
R
R
HR

IS 1134C (SC 203)

India

Durra

HR

IS 1139C (SC 205)

India

Durra

HR

IS 1144C (SC 451)

India

Durra

HR

Ax, Ab

IS 1366C (SC 210)

India

Durra

Ax

IS 1598C (SC 214)

India

Dochna

HR

Ax, Ab

IS
IS
IS
IS
IS

1840
2312
3796
4657
5188C (SC 245)

USA
USA
India
India

Durra
Caffrorum
Durra
Durra/Dochna

R
R
MR
R
HR

Ab

IS 5490
IS 5887C (SC 248)

India
India

Durra
Roxburghii

R
HR

Ax

IS 6389C (SC 489)

India

Nandyal

HR

IS 6416C

India

Nandyal

HR

Ax

IS 6426C (SC 497)

India

Nandyal

HR

Ax

IS 6962C

India

Caudatum

IS 8100C

Japan

HR

IS 12158C (SC 984)

Ethiopia

Caudatum/
Nigricans
Bicolor

HR

Ax, Ab

IS 12551C (SC 31)

Ethiopia

Caudatum

HR

Ax, Ab

IS 12599C (SC 90)

Congo

Guinea

HR

Ax, Ab

IS 12608C (PI 257 595,

SC 108-14-E)

Ethiopia

Caudatum

HR

Ax, Ab

IS 12609C (SC 109)

Ethiopia

Caudatum

IS 12610C (SC 110-14)

Ethiopia

Caudatum

IS 12611C (SC 111 or

SC 170)

Ethiopia

Bicolor

Zimbabwe
Botswana,
Zimbabwe
Botswana,
Taiwan
Zimbabwe
Botswana,

IS 12612C

Ethiopia

Bicolor

Zimbabwe
Japan,

IS 12637C (SC 146)

Ethiopia

Bicolor

Ab

IS 12645C

Ethiopia

Bicolor

HR

Ax, Ab

South Africa
Botswana,
Zimbabwe
Botswana,
Zimbabwe

Teetes
Teetes
Teetes
Teetes

et
et
et
et

al.
al.
al.
al.

(1995)
(1995)
(1995)
(1995)

Teetes et al. (1995)

Teetes et al. (1995)

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745

Table 3 (continued)
Germplasm accession

IS 12661C

Origin

Ethiopia

Classication

Bicolor

Resistance
Level

Mechanism

HR

Ax, Ab

Country from
which reported

Reference

Botswana,

Manthe et al. (1984), Manthe (1992), and

Teetes et al. (1995)
Manthe et al. (1984), Manthe (1992), and
Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992) and Teetes et al. (1995)
Manthe (1992)
Manthe (1992)
Balikai (1993)
Peterson (2002)
Balikai (1993)

Zimbabwe
IS 12664C

Ethiopia

Bicolor

HR

IS 12667C

Ethiopia

Bicolor

IS 14048
IS 23250 (Sima)

Malawi

Guinea
Caudatum
Bicolor
Durra

R
R

Botswana,
Zimbabwe
Botswana,
Zimbabwe
India
South Africa

India

IS 33843

Ax, Ab

Ax, antixenosis; Ab, antibiosis; HR, highly resistant; MR, moderately resistant; and R, resistant.

although resistant sources served as suitable hosts in

connement studies, which indicate the involvement of
both antixenosis and antibiosis mechanisms. Antixenosis for adult colonization was noticed in TAM 428, IS
1144C, IS 1366C, IS 1598C, IS 6416C, IS 6426C, IS
12661C, and IS 12664C. Among them, IS 1144C and IS
12664C were preferred less than the resistant check,
TAM 428. High levels of antibiosis expressed in TAM
428, IS 1144C, IS 5188C, IS 12609C, and IS 12664C for
least number of days to reproduction; TAM 428, IS
12609C, and IS 12664C for greater mortality and
shorter longevity of adults, and production of fewer or
no nymphs (Teetes et al., 1995).
Morpho-physiological traits: In sorghum, some of the
morpho-physiological traits such as genotypes with
small, narrow, and fewer leaves, and low leaf
bending at the seedling stage (Mote and Kadam,
1984), greater plant height and greater distance between
two leaves and the presence of waxy lamina (Mote and
Shahane, 1994), and epicuticular wax on the ventral
surface of the leaves were associated with reduced
susceptibility to the sugarcane aphid (Pi and Hsieh,
1982a, b).
Biochemical factors: Host suitability to various
phloem-feeding Homoptera has frequently been related
to nitrogen levels in host plants. Hsieh (1988) stated that
the presence of p-hydroxybenzaldehyde during HCN
release from the sorghum leaves due to aphid biting, at
the seedling stage, may be important to repel further
attack. The mean HCN content of F1 hybrids produced
from crosses between high and low HCN content
genotypes usually had correlations intermediate between
those of the parents or lines were closer to the parent
with a high HCN content. The development of aphid
populations and leaf sugary exudation was more
pronounced in sorghum genotypes having higher nitro-

gen, sugar, and chlorophyll content of leaves (Mote and

Jadhav, 1993; Mote and Shahane, 1994). The varieties
ICSV 9, BTP 28, IS 1640, ICSV 148, and Swati (SPV
504) with higher contents of phosphorus, potassium,
and polyphenols were less preferred by the sugarcane
aphids and also showed less development of leaf sugary
exudation (Mote and Shahane, 1994).
There is a signicant reduction in nitrogen, phosphorus, potash, total sugars, and chlorophyll content in
sorghum due to infestation by the sugarcane aphid. In
contrast, there is a signicant reduction in polyphenols
in resistant over the susceptible sorghum genotypes
(Balikai, 2001). In India, planting in the third week of
September caused a reduction in protein, total minerals,
and fat content in the grain due to infestation by the
sugarcane aphid. Similarly, there is also a decline in
diastase activity in sorghum grain, while there is an
increase in crude ber and carbohydrate content when
infested by M. sacchari (van den Berg et al., 2003). In
sorghum fodder, there was a loss of 10% and 7.0% in
crude protein and crude ber content, respectively
(Balikai, 2001).
Genetic basis of resistance: Greenhouse and eld
studies with the crosses between PI 257595 (highly
resistant), 129-3A (moderately resistant), and RTx 430
(susceptible) have shown that resistance is monogenic
and controlled by a single dominant gene (Hsieh and Pi,
1982; Pi and Hsieh, 1982b; Tan et al., 1985). Studies
have also indicated that PI 257595 and 129-3A have the
same gene for resistance, although the resistance gene of
129-3A has modiers (Pi and Hsieh, 1982a). Although
dominant and additive gene actions are involved,
additive gene action accounts for the resistance expression (Hsieh and Pi, 1988). The cross between RTx
430  129-3A indicated the presence of complimentary
gene action (Chang and Fang, 1984).

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Table 4
Parental lines (A/B- and R-lines) of sorghum resistant to the sugarcane aphid, M. sacchari (Zehnt.) reported from different countries
Parental line

Resistance
Level

Country from which reported

Reference

Mechanism

A/B-line
2B
9B
27A
33B
42B

R
R
R
R
R

India
India
India
India
India

53B
104A/B
116A/B
117B
129-3A

R
R
R
R
R

205B
296B
2077A/B
2219A/B
4692A
A 36227B
AB 31
AKMS 14A
ICSB 53
ICSB 70
ICSB 84
ICSB 90
ICSB 101
ICSB 2730
ICSB 2731
ICSB 88001
ICSB 88014

R
R
R
R
MR
R
MR
R
R
R
R
R
R
HR
HR
R
R

India
India
India
India
Taiwan,
South Africa
India
India
India
India
Japan
India
China
India
India
India
India
India
India
India
India
India
India

Balikai (2001)
Balikai (2001)
AICSIP (1998)
Patil (1992)
AICSIP (1990, 1998), Patil (1992), and
Balikai (1993)
Patil (1992)
AICSIP (1998) and Balikai (2001)
AICSIP (1991, 1998) and Patil (1992)
Patil (1992) and AICSIP (1996)
Pi and Hsieh (1982a)
Wenzell et al. (1998)
Patil (1992) and AICSIP (1996)
Kishore (2000)
Balikai (2001)
Patil (1992) and Balikai (1993)
Hagio and Ono (1986)
AICSIP (1991)
Chang (1981a, b)
AICSIP (1997)
AICSIP (1991)
Patil (1992)
Patil (1992)
Patil (1992)
Patil (1992)
AICSIP (2003)
AICSIP (2003)
AICSIP (1991)
AICSIP (1991)

R-line
2R

5R

China,
South Africa
China

Chang (1981a, b)
Wenzell et al. (1998)
Chang (1981a, b)

47R
57R
C 43
C 81
ICSR 160
ICSR 161
ICSR 162
ICSR 165
ICSR 172
ICSR 174
ICSR 194
ICSR 38111
ICSR 89008
ICSR 89009
M 148-138
R 128
R 131
R 132
R 1413
R 354
RS 29

MR
MR
HR
R
R
R
R
R
R
R
R
HR
R
R
R
R
R
R
R
R
R

China
China
India
India
India
India
India
India
India
India
India
India
India
India
India
China
China
China
India
India
India

RS 67

India

RS 291
RS 530
RS 649
RTAM 428

R
R
R
HR

SB 1085

India
India
India
China,
South Africa
India

Chang (1981a, b)
Chang (1981a, b)
AICSIP (2003)
Balikai (1993)
AICSIP (1991)
Patil (1992)
Patil (1992)
AICSIP (1991) and Patil (1992)
Patil (1992)
Patil (1992)
AICSIP (1991)
AICSIP (2003)
AICSIP (1991)
AICSIP (1991)
AICSIP (1990)
Chang and Fang (1984)
Chang and Fang (1984)
Chang and Fang (1984)
Balikai (1993)
Kishore (2000)
Patil (1992), Balikai (1993),
Kishore (2000), and AICSIP (2003)
Patil (1992), Balikai (1993),
Kishore (2000), and AICSIP (2003)
AICSIP (1997)
AICSIP (1989)
AICSIP (1998)
Xu (1982)
Wenzell et al. (1998)
AICSIP (1997)

Ax, Ab
Ax, Ab

Ax, Ab

Ax, Ab

Ax, Ab

Ax, Ab

Ax, antixenosis; Ab, antibiosis; HR, highly resistant; MR, moderately resistant; and R, resistant.

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Table 5
Agronomic elite lines/experimental varieties of sorghum resistant to the sugarcane aphid, M. sacchari (Zehnt.) reported from different countries
Agronomic elite line/
experimental variety

Resistance
Level

Country from
which reported

Reference

Taiwan
China
India
South Africa
India
India
India
South Africa
South Africa
South Africa
India
China
India
India
India
India
India
India
India
India
India
India
India
India
India
India
India
Japan
India
Japan

Pi and Hsieh ( 1982a)

Wang et al. (1991)
Mote and Shahane (1988, 1993)
Peterson (2002)
AICSIP (1982)
AICSIP (1982)
AICSIP (1980)
Peterson (2002)
Peterson (2002)
Peterson (2002)
Mote et al. (1985)
Chang (1981a, b)
Mote and Shahane (1988, 1993)
AICSIP (1988) and Mote and Shahane (1988, 1993)
Sharma (1993)
Sharma (1993)
Sharma (1993)
AICSIP (1991)
AICSIP (1998)
AICSIP (1998)
AICSIP (1998)
AICSIP (1999)
AICSIP (2000)
AICSIP (2000)
AICSIP (2000)
AICSIP (2000)
AICSIP (2000)
Hagio and Ono (1986)
AICSIP (1990)
Hagio et al. (1985), Hagio and Ono (1986), and
Hagio (1992)
Teetes et al. (1995)
Teetes et al. (1995)
Patil (1992)
Wenzell et al. (1998)
Peterson (2002)
AICSIP (1980)
AICSIP (1982)
AICSIP (1982)
AICSIP (1980, 1982)
AICSIP (1982)
AICSIP (1982)
AICSIP (1982)
AICSIP (1982)
AICSIP (1982)
AICSIP (1982)
AICSIP (1980)
AICSIP (1982)
AICSIP (1982)
Mote and Kadam, 1984)
AICSIP (1997)
Mote and Shahane (1988, 1993)
Mote and Kadam, 1984)
Patil (1992)
Patil (1992)
Patil (1992)
Patil (1992)
Patil (1992)
Patil (1992)
Patil (1992)

Mechanism

Agr 4S2560 Shallu

7511
BPT 28
CE 151
E 108
E 119
E 303
Ent. 62SADC
FGYQ 336
FGYQ 353
Hami
HB 37
ICSV 9
ICSV 148
ICSV 197
ICSV 743
ICSV 745
ICSV 89013
ICSV 93028
ICSV 93038
ICSV 93093
ICSV 95081
ICSV 95085
ICSV 95093
ICSV 95099
ICSV 95110
ICSV 96081
NK 266
NR 349
PE 954 177

HR
HR
R
R
R
R
R
R
R
R
R
R
R
R
R
R
R
R
R
R
R
R
R
R
R
R
R
MR
R
HR

Ax, Ab

PNR 8537

HR

Ax

PVR 10
SA 967
SDSL 89426
SPV 97
SPV 101
SPV 102
SPV 224
SPV 232
SPV 236
SPV 243
SPV 245
SPV 257
SPV 291
SPV 303
SPV 378
SPV 386
SPV 422
SPV 492
SPV 504
SPV 625
SPV 969
SPV 991
SPV 998
SPV 1002
SPV 1004
SPV 1051
SPV 1053

R
R
R
R
R
R
R
R
R
R
R
R
R
R
R
R
MR
R
R
MR
R
R
R
R
R
R
R

Ax, Ab

Botswana,
Zimbabwe
India
South Africa
South Africa
India
India
India
India
India
India
India
India
India
India
India
India
India
India
India
India
India
India
India
India
India
India
India
India

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Table 5 (continued)
Agronomic elite line/
experimental variety

Resistance
Level

SPV
SPV
SPV
SPV
SPV
SPV
SPV
SPV
SPV
SPV
SPV

1054
1155
1178
1359
1380
1413
1450
1452
1453
1462
1465

R
R
R
R
R
R
R
R
R
R
R

Country from
which reported

Reference

India
India
India
India
India
India
India
India
India
India
India

Patil (1992)
Kishore (2000)
Balikai (1993)
Kishore (2000)
Kishore (2000)
Kishore (2000)
Kishore (2000)
Kishore (2000)
Kishore (2000)
Kishore (2000)
Kishore (2000)

Mechanism

Ax, antixenosis; Ab, antibiosis; HR, highly resistant; MR, moderately resistant; and R, resistant.

8. Virus vector

10. Natural enemies

The sugarcane aphid is a vector and transmits three

persistent viruses namely, millet red leaf virus (Blackman and Eastop, 1984), and sugarcane yellow leaf virus
on sorghum and sugarcane (Schenck, 2000), and
sugarcane mosaic virus (SMV) on sorghum (Bhargava
et al., 1971; Kondaiah and Nayudu, 1984; Setokuchi
and Muta, 1993). Yang (1986) demonstrated that both
the sugarcane aphid and corn leaf aphid (Rhopalosiphum
maidis) were efcient transmitters of SMV between
sorghum, corn, and sugarcane. Latent periods of 30 and
20 days were observed for SMV on corn and sweet
sorghum, respectively. However, the disease symptoms
colonized by M. sacchari takes a longer time to express
on sweet sorghum than on corn or sugarcane (Yang,
1986).

Over 47 species of natural enemies attack M. sacchari

worldwide. Although unable to prevent the buildup of
damaging numbers of the sugarcane aphid even in the
season, natural enemies play a very important role (van
Rensburg and van Hamburg, 1975), and often maintain
the sugarcane aphid populations below the economic
threshold levels (ETLs) in sorghum (van Rensburg,
1973b; Anonymous, 1978; Chang, 1981a, b; Meksongsee
and Chawanapong, 1985) (Table 7).
Zimmerman (1948) reported Aphelinus maidis parasitizing the sugarcane aphid in Hawaii. Similarly, the
parasitoids Enrischia comperei Ashm. in Australia (Gilstrap, 1980), Exochonus concavus (Fursch), Leucopus
spp., and Lioadalia flavomaculata (DeGeer) in South
Africa (van Rensburg, 1973b) have been recorded. The
aphid parasite, Lysiphlebus testaceipes (Cresson) (Hymenoptera: Braconidae) attacks the sugarcane aphid in
Hawaii (Zimmerman, 1948) and indigenous L. dehliensis
Zehntner is credited with biological control on sugarcane in India (Varma et al., 1978). Low levels of
parasitism by unidentied parasitoids have been observed in sorghum in South Africa, but generally are
considered less important in suppressing the aphid
population abundance (Anonymous, 1978; van Rensburg, 1973b).
Emphasis has been placed on predators, primarily
ladybeetles (Coleoptera: Coccinellidae), lacewings (Neuroptera: Chrysomelidae and Hemerobiidae), and hoveries (Diptera: Syrphidae) as species causing greatest
mortality to the sugarcane aphid populations. Numerous predators suppressed sugarcane aphid abundance
on sorghum in South Africa (Anonymous, 1978; van
Rensburg, 1973a, b), Tanzania (Bohlen, 1973), and
Sudan (Schmutterer, 1969). In southern Africa, important predator species are ladybeetles, Cheilomenes
propinqua (Muls.) var. quadrilineata (Muls.), C. lunata
(Fabr.), C. sulphurea (Fabr.), L. flavomaculata (De Geer),

9. Cultural practices
Among the cultural practices, early planting results in
the crop escaping from aphid attack (van Rensburg,
1974) and high plant density promotes low plant vigor
and concomitantly reduces aphid abundance (Flattery,
1982; van Rensburg, 1979; Setokuchi, 1975). Cutting of
forage sorghum before the rst week of aphid abundance prevents not only the damage but also regulates
subsequent increase in aphid density on a ratoon crop
(Setokuchi, 1977). Since the sugarcane aphid overwinters on ratoon sorghum, S. verticilliflorum, S.
halepense, P. maximum, and Setaria spp., their destruction before the sorghum crop is planted reduces the
carryover of the pest. Mulching with rice/wheat straw is
another practice effective in reducing colonization by
aphids. Sanchez and Cermeli (1987) used yellow traps
with water to capture the migrant aphids in the elds of
sorghum to forecast their migratory pattern and
population dynamics in Venezuela.

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Table 6
Hybrids, varieties, and locals of sorghum resistant to the sugarcane aphid, M. sacchari (Zehnt.) reported from different countries
Hybrid/variety/local

Resistance

Country from
which reported

Reference

R
R
R
R
R
R
R

South
South
South
South
South
South
South

Wenzell et al. (1998)

Wenzell et al. (1998)
Wenzell et al. (1998)
Wenzell et al. (1998)
Wenzell et al. (1998)
Wenzell et al. (1998)
Peterson (2002)

R
R
R

South Africa
South Africa
South Africa

Peterson (2002)
Peterson (2002)
Peterson (2002)

R
R
HR
R

South Africa
South Africa
India
India

Wang et al. (1991)

Wang et al. (1991)
Balikai (1993)
AICSIP (1996)

India
India

Balikai (1993, 2001)

Kishore (2000)

Hagio (1987)
Xu (1982)
Mote et al. (1985)
Hagio (1967)
AICSIP (1982, 1987)
Hagio (1987)
Hagio (1987)
Mote et al. (1985)
Hagio (1992)
Mote et al. (1985)
Mote et al. (1985)
Teetes et al. (1995)
Teetes et al. (1995)
Hagio (1987)
Manthe et al. (1992)
Chang (1981a, b)
Mote et al. (1985)
Setokuchi (1976), Hagio et al. (1985),
Hagio and Ono (1986), and Hagio (1992)
Hagio and Ono (1986)
Hagio (1987)
Hagio (1992)
Hagio et al. (1985), Hagio and Ono (1986), and
Hagio (1992)

Level
Hybrid
272 x 3122
846 x 1469
1158 x 132
1175 x 551
1470 derivative
1670 x 1110
(6BRON161/(7E0366Tx2783)HG54)CE151)-CG-3-BGBK
(CE1518BDM499)-LD17-BD1
(MaciaTAM 428)-1-1-2
(MR112B-92M2Tx 2880)-SM3SM1-ML 52
7511 x Kang 60-P341
7511 x Kang60-P400
CSH 13K&R
SPH 634

Mechanism

Ax, Ab

Africa
Africa
Africa
Africa
Africa
Africa
Africa

Variety
CSV 8R
Sel 3

R
R

Local
Boushi Kaoliang
Dabor
Dagadi
Dairyukoku
DJ 6514
Gyushinpaku
Hakubai Kaoliang
Jogri
Koumairou
Kuchkuchi
Lakadi
Ludende

R
HR
R
R
R
R
R
R
HR
R
R
HR

Minasel
SA 1429
SC 110-14
PJ 4R
Senkinshiro

R
R
R
R
HR

Japan
China
India
Japan
India
Japan
Japan
India
Japan
India
India
Botswana,
Zimbabwe
Japan
South Africa
Taiwan
India
Japan

Setokho-2-gou
Shoukoubai
Shoumai Kaoliang
Suzuho

MR
R
HR
HR

Japan
Japan
Japan
Japan

Ax, Ab

Ax, Ab

Ax, Ab

Ax, Ab
Ax, Ab

Ax, antixenosis; Ab, antibiosis; HR, highly resistant; MR, moderately resistant; and R, resistant.

E. concavus (Furch), Chilocorus nigritus (F.), and

Scymnus morelleti (Muls.) (Coleoptera: Coccinellidae)
in South Africa (van Rensburg, 1973b), S. babar Sassji
in China (He et al., 1987), and Diomus terminatus (Say)
in the USA (Hall, 1987). He et al. (1987) also reported
that adults of S. babar overwintered in the leaf sheaths
of Juncellus serotinus, Acorus calamus L., Typha
latifolia, and Boiboschnoenus mantinus. They become

active in mid-April and migrated to the sorghum elds

in mid-June to predate upon M. sacchari and then into
maize elds in early August to feed on R. maidis and R.
padi, and complete one and two generations on sorghum
and maize, respectively.
Among the dipteran predators, syrphids also play an
important role on the sugarcane aphid viz., Allograpta
exotica (Wiedemann, 1830) in the USA (Florida)

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Table 7
Natural enemies of the sugarcane aphid, M. sacchari (Zehnt.) reported in different countries
Natural enemy

Order

Family

Country

Reference

Pathogen
Verticillium lecanii (Zimm.) Viegas

Hypocreales

Hypocreaceae

USA (Florida)

Hall (1987)

Parasitoid
Aphelinus maidis Timberlake
Enrischia comperei ashm.
Bracon sp.
Lioadalia flavomaculata (DeGeer)
Lysiphlebus dehliensis Zehntner
Lysiphlebus testaceipes (Cresson)

Hymenoptera
Hymenoptera
Hymenoptera
Hymenoptera
Hymenoptera
Hymenoptera

Aphelinidae
Elasmidae
Braconidae
Aphelinidae
Braconidae
Braconidae

USA (Hawaii)
Australia
India
South Africa
India
USA (Florida),
USA (Hawaii)

Zimmerman (1948) and Gilstrap (1980)

Gilstrap (1980)
Hussain (1991)
van Rensburg (1973)
Varma et al. (1978)
Hall (1987)
Zimmerman (1948)

Diptera
Coleoptera

Syrphidae
Coccinellidae

USA (Florida)
India

Hall (1987)
Seshu Reddy and Davies (1979)

Coleoptera
Coleoptera

Coccinellidae
Coccinellidae

Chrysoperla sp.
Chrysoperla basalis Walk.
Chrysoperla collaris Schneider
Chrysoperla externa (Hagan)
Coelophora inaequalis (Fabricius)

Neuroptera
Neuroptera
Neuroptera
Neuroptera
Coleoptera

Chrysomelidae
Chrysomelidae
Chrysomelidae
Chrysomelidae
Coccinellidae

Coleomegilla maculata Deg.

Coleomegilla maculata fuscilabris
(Mulsant)
Cycloneda sanguinea (L.)

Coleoptera
Coleoptera

Coccinellidae
Coccinellidae

Puerto Rico
India,
South Africa
India
Thailand
Puerto Rico
USA (Florida)
Australia,
USA (Hawaii)
Dutch Guiana
USA (Florida)

Gilstrap (1980)
Gilstrap (1980)
van Rensburg (1973)
Seshu Reddy and Davies (1979)
Meksongsee and Chawanapong (1985)
Gilstrap (1980)
Hall (1988)
Gilstrap (1980)
Gilstrap (1980)
Gilstrap (1980)
Hall (1988)

Coleoptera

Coccinellidae

Diomus terminatus Say

Exochomus concavus (Furch)
Geocoris sp.
Hippodamia convergens Guerin
Illeis indica Timberlake
Leucopus spp.
Menochilus (Cheilomenes) lunata
(Fabr.)
Menochilus (Cheilomenes) propinqua
(Muls.) var quadrilineata
Menochilus (Cheilomenes) sexmaculatus
(Schall.)

Coleoptera
Coleoptera
Hemiptera
Coleoptera
Coleoptera
Diptera
Coleoptera

Coccinellidae
Coccinellidae
Lygaeidae
Coccinellidae
Coccinellidae
Chamaemyiidae
Coccinellidae

Dutch Guiana,
Puerto Rico,
USA (Florida)
USA (Florida)
South Africa
India
USA (Florida)
India
South Africa
South Africa

Gilstrap (1980)
Gilstrap (1980)
Hall (1988)
Hall (1987)
van Rensburg (1973)
Seshu Reddy and Davies (1979)
Hall (1988)
Seshu Reddy and Davies (1979)
van Rensburg (1973b)
van Rensburg (1973b)

Coleoptera

Coccinellidae

South Africa

van Rensburg (1973b)

Coleoptera

Coccinellidae

India,

Menochilus (Cheilomenes) sulphurea

(Fabr.)
Micraspis discolor (Fabr.)
Micromus subanticus (Walker)
Nacoleia (Omiodes) accepta Btlr.
Ola v-nigrum Mulsant
Phaenobremiameridionalis Felt.
Prospaltella transvena Timb.
Scoloposcelis parallelus Motsch.
Scymnodes (Platynomus) lividigaster
(Mulsant)
Scymnus babar Sassji
Scymnus loewi Muls.
Scymnus morelletti (Muls.)
Scymnus nubilus Mulsant
Scymnus roseicollis Muls.
Syrphus balteatus (DeGeer)
Xanthogramma aegyptium (Wied.)

Coleoptera

Coccinellidae

Thailand
South Africa

Seshu Reddy and Davies (1979) and

Gilstrap (1980)
Meksongsee and Chawanapong (1985)
van Rensburg (1976)

Coleoptera
Neuroptera
Diptera
Coleoptera
Diptera
Hymenoptera
Hemiptera
Coleoptera

Coccinellidae
Hemerobiidae
Cecidomyiidae
Coccinellidae
Cecidomyiidae
Aphelinidae
Anthocoridae
Coccinellidae

Thailand
USA (Florida)
USA (Hawaii)
USA (Florida)
USA (Hawaii)
USA (Hawaii)
Puerto Rico
Australia

Meksongsee and Chawanapong (1985)

Hall (1988)
Gilstrap (1980)
Hall (1988)
Gilstrap (1980)
Zimmerman (1948) and Gilstrap (1980)
Gilstrap (1980)
Gilstrap (1980)

Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Diptera
Diptera

Coccinellidae
Coccinellidae
Coccinellidae
Coccinellidae
Coccinellidae
Syrphidae
Syrphidae

Diptera

Syrphidae

China
Puerto Rico
South Africa
India
Puerto Rico
Thailand
USA (Louisiana)
South Africa
India

He et al. (1987)
Gilstrap (1980)
van Rensburg (1976)
Gilstrap (1980)
Gilstrap (1980)
Meksongsee and Chawanapong (1985)
White et al. (2001)
van Rensburg (1976)
Seshu Reddy and Davies (1979)

Predator
Allograpta exotica (Wiedemann, 1830)
Brumus suturalis (Fabr.)
(Syn: Brumoides suturalis (Fabr.))
Ceratomegilla (Megilla) inonata Muls.
Chilochorus nigritus (F.)

Xanthogramma scutellaris

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(Hall, 1987), Syrphus balteatus (DeGeer) in Thailand

(Meksongsee and Chawanapong, 1985), Xanthogramma
aegyptium in the USA (Louisiana) (White, 1987), South
Africa (van Rensburg, 1973a, b), and X. scutellarius in
India (Seshu Reddy and Davies, 1979); and Leucopus
spp. (Chamaemyiidae) in South Africa (van Rensburg,
1973b). Hall (1988) reported Micromus subanticus
(Walker) (Neuroptera: Hemerobiidae) as feeding on
the sugarcane aphid in the USA (Florida). Other
chrysopids have also been recorded such as Chrysoperla
basalis Walker, C. collaris Schneider, and C. externa
(Hagan) in Thailand (Meksongsee and Chawanapong,
1985), Puerto Rico (Gilstrap, 1980) and the USA
(Florida) (Hall, 1988).
Hall (1987) found that the entomogenous fungus,
Verticillium lecanii (Zimm.) Viegas is an important
biological control agent in the USA (Florida). However,
ants are notorious in interfering with the benecial
activities of aphid predators and/or parasites and the
sugarcane aphid may sometimes benet from a symbiotic association with certain ant species.

11. Chemical control

Soil application of systemic insecticides such as
disulfoton, disyston, and phorate (van Rensburg and
van Hamburg, 1976; Mote, 1977; Denmark, 1988) were
effective, with phorate superior to disulfoton (Mote,
1977). Foliar sprays of demeton-S-methyl, dimethoate,
endosulfan, and parathion (van Rensburg, 1979; Mote,
1985); diazinon, malathion, metasystox, and phosdrin
(Denmark, 1988); quinalphos (Chaudhari et al., 1994);
and carbofenthion and carbofuran were effective when
the populations reach 70 and 155 aphids plant 1 at 50
and 80 days after planting in spring and 60 and 90 days
after planting in autumn, respectively (Fang, 1990).
Spraying extracts of leaves and kernels of Vinca rosea,
Pongamia pinnata, Azadirachta indica, and Vitex negundo took 10 days to cause mortality of aphids:
Dimethoate, chlorpyriphos, endosulfan, alfamectin,
and malathion not only reduced aphid populations but
promoted grain and fodder yields, besides increase in
kernel weight (Balikai, 2001). Currently, the only
effective method to control the M. sacchari populations
is the use of insecticides, but thus use is not considered
economically feasible for small-scale farmers in developing countries (van Rensburg and van Hamburg, 1975;
Young and Teetes, 1977; Page et al., 1985). This is
because, in the absence of natural enemies, aphid
infestations may resurge, so a second or even a third
insecticide application may be required for control (van
Rensburg, 1978, 1979). Systemic insecticides are effective against the sugarcane aphid, but do not kill the
benecial arthropods (van Rensburg et al., 1978; van
Rensburg, 1980). However, application of demeton-S-

751

methyl, parathion, and monocrotophos resulted in

mortality of coccinellids and syrphids while endosulfan
and pirimicarb were less toxic to predators (van
Rensburg and van Hamburg, 1975).

12. Future research

Several factors affect the plant responses to M.
sacchari injury such as the time of infestation (season
and plant growth stage), duration of the infestation,
environmental stresses (especially drought) and nutritional status of the host plant, which cumulatively affect
the yield loss. The sugarcane aphid feeding at the basal
leaves had a strong inuence on mobility of mineral
nutrients, amino compounds, and carbohydrates in the
phloem. By feeding at these sites, M. sacchari alters the
carbohydrate-partitioning patterns of sorghum, suggesting that infestation might alter sink-source relationships
within the plant. Very few studies have investigated the
impact of environmental factors on M. sacchari damage
symptom development, which mostly depends on the
initial aphid density colonized as well as the inherent
plant resistance. Thus, the impact of abiotic factors such
as temperature and photoperiod on the life history traits
of M. sacchari needs to be investigated. Further, the
contribution of weeds and the native vegetation to the
sugarcane aphid population dynamics is not clearly
known, although considered as a major factor inuencing sorghum infestation. Diversity of natural enemies
may vary within and among cultivar communities
depending on a variety of factors including microclimate, prey density, annual changes in host plant species
abundance, the timing and rate of migration and
dispersal, and geographic location. The epidemiology
of transmission of three persistent viruses transmitted by
the sugarcane aphid has not been substantially investigated. Emphasis should be given on the population
colonization at the seedling stage of sorghum because
they may have greater impact affecting the growth and
development of the crop.
The impact of natural enemies on sugarcane aphid
populations in sorghum is not well understood. Quantication and predicting the impact of natural enemies, as
a group or as individuals, has been difcult because of
the many species involved. The natural enemies have a
greater effect early in the season, since the increase per
generation of the sugarcane aphid population early in
the season is slow compared with late season increases.
Comprehensive information needs to be generated on
the population dynamics of M. sacchari and the factors
that lead to rapid population buildup. Environmental
factors are largely responsible for the initial decline in
aphid levels following a summer outbreak (Hall, 1987).
Quantication of development rates of immatures, and
potential population growth rates of the sugarcane

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aphid under different temperature regimes similar to

those occurring in the post-rainy season sorghum niche
in India would greatly aid in forecasting models.
Drought and sugarcane aphid infestation are two
factors that individually cause yield reduction in
sorghum, but when they occur in succession results in
a synergistic interaction between aphid infestation and
predisposing the crop to drought stress. Field observations have indicated that aphid damage is greater when
drought also occurs as a stress factor. The magnitude of
plant damage to sorghum in relation to the density of
M. sacchari as well as the stage of plant growth is not
known. It may be that sorghum can recover and
compensate for damage due to M. sacchari as has been
observed by the feeding injury caused by S. flava in
sorghum. Insect density relationships with yield loss are
not available. It is not known, whether the sugarcane
aphid injects a phytotoxin during feeding that degrades
chloroplasts or the damage results from the plants
response to mechanical injury. As little is known about
how aphids elicit these changes and the physiological
mechanisms associated with injury, improving our
understanding of how M. sacchari affects the sorghum
plant physiology will be important in identifying new
targets for aphid resistance in sorghum.
Cultural practices such as elimination of volunteer
sorghum plants from the previous crop and the
destruction of susceptible weedy hosts needs greater
emphasis. Search for new sources of resistance is
essential to diversify the genetic background. Identication of resistant sources in elite background as well as in
parental lines in the development of hybrids, and
knowledge on the mechanisms would greatly assist in
developing cultivars with stable sources of resistance to
the sugarcane aphid, under high yield background.
Combinations of different categories have effects that
may be more benecial and compatible in trophic
interactions than the effect of individual components
of resistance.

Acknowledgements
We are greatly indebted to Dr. Mangala Rai, Director
General, Indian Council of Agricultural Research, New
Delhi for his constant encouragement and keen interest.

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