27

1Texas

IPM Case Studies: Sorghum

Gerald J. Michels, Jr.1 and John D. Burd2

Agricultural Experiment Station, Bushland, TX 79012, USA; 2Plant Science and Water Conservation Laboratory, USDA-ARS, Stillwater, OK 74075, USA

Introduction
Sorghum, Sorghum bicolour (Fig. 27.1), also known as great millet and Guinea corn, originated in Africa and is cultivated throughout the tropical, subtropical, and warm temperate areas of the world. Sorghum is grown for animal feed and forage, human consumption, and for fibre (FAO, 1979; Dibb, 1983). Worldwide, 41.5 million ha of sorghum were harvested in 2001 (FAO, 2001). There are four important worldwide aphid pests of sorghum: Schizaphis graminum (greenbug), Rhopalosiphum maidis (corn leaf aphid), Melanaphis sorghi (sugarcane aphid), and Sipha flava (yellow sugarcane aphid) (Young and Teetes, 1977). S. graminum is the key cosmopolitan aphid pest of sorghum listed by Young and Teetes (1977). The other three species are considered occasional pests. There is an extensive literature base on IPM components for the greenbug on sorghum. However, few studies have compared integrated and single-method management systems, including an economic analysis. Due to the startling lack of IPM case studies addressing other sorghum-feeding aphids, this review will concentrate on greenbug IPM practices, especially since greenbug has been studied for many years and has a cosmopolitan distribution. It is hoped that this review will
CAB International 2007. Aphids as Crop Pests (eds H. van Emden and R. Harrington)

stimulate research comparing single and multiple-component control strategies for aphid pests of sorghum.

A Short History of Schizaphis graminum on Sorghum

Schizaphis graminum (Figs 27.2 and 27.3) has been a key pest of sorghum for a relatively short period of time. Greenbugs were known to utilize sorghum as a host as early as 1863 in Italy (Hunter, 1909) and were noted on sorghum in Africa by Matthee (1962), and again in Europe by Barbulescu (1964). The greenbug was first described as a significant pest of sorghum in the USA in 1968 (USDA, 1968) when serious damage to sorghum was reported in the southwestern and Great Plains regions. This outbreak resulted in millions of hectares being treated with insecticides to control the pest. Harvey and Hackerott (1969) reported that over 400,000 ha of sorghum in Kansas were infested with greenbugs and that the pest destroyed over 100,000 ha of the crop. Harvey and Hackerott (1969) designated this sorghum-damaging greenbug as biotype C, based on its ability to damage Piper Sudan grass (Sorghum drummondii), which was resistant to biotype B greenbugs.

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Fig. 27.1. Healthy, irrigated grain sorghum at the Texas Agricultural Experiment Station North Plains Research Field, Etter, Texas (photo by G.J. Michels,

Fig. 27.2.

The greenbug, Schizaphis graminum (original colour photo courtesy of R.W. Behle).

By 1981, greenbugs were estimated to be responsible for 2.5% of the 9.0% annual loss in grain sorghum due to insects, and it ranked as the second most damaging insect

pest of sorghum in the USA (USDA, 1981). As late as 1992, 31% of sorghum in the USA was treated with insecticides to control greenbugs (Webster et al., 1995).

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Fig. 27.3. Greenbug damage to grain sorghum at the Texas Agricultural Experiment Station North Plains Research Field, Etter, Texas (photo by G.J. Michels, Jr.).

Current Greenbug Management Practices

In 1981, the US Department of Agriculture (USDA, 1981) stated: The controls for reducing greenbug damage are as complete as those for any insect. At the time of the report, IPM components, especially resistant hybrids, were available for use by producers.

Chemical control Research examining the efficacy of foliar and soil-applied insecticides to control greenbug began in 1968 and articles appeared in the early 1970s (DePew, 1971, 1974; Cate et al., 1973a,b). Foliar organophosphates, carbamates, and even one organochlorine (endrin) gave excellent control (DePew, 1971; Cate et al.,

1973a). Soil-applied organophosphates also provided control, while soil-applied carbamates varied in their efficacy (DePew, 1974). Within a few years, however, greenbug resistance to organophosphates, specifically disulfoton, was reported (Peters et al., 1975; Teetes et al., 1975). Additional organophosphates and carbamates were developed throughout the 1970s and 1980s, most notably chlorpyrifos, which became the chemical of choice in the 1980s. However, greenbug resistance to chlorpyrifos has been documented (Niemczyk and Moser, 1982; Sloderbeck et al., 1991). Research conducted by Shufran et al. (1996, 1997a,b) and Rider et al. (1998) showed that greenbug genotypes differed in their levels of insecticide resistance. Shufran et al. (1997b) and Stone et al. (2000) demonstrated that life history

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parameters of these genotypes differed, with the most prevalent insecticide-resistant one having none of the reproductive disadvantages often associated with insecticideresistant arthropods. Archer et al. (1999) concluded that combinations of chlorpyrifos and malathion applied to a 3 : 1 mixture of insecticide-resistant and -susceptible greenbug populations resulted in a significant net return when compared to untreated sorghum, or even when the greenbugs received a pretreatment application of chlorpyrifos, which supposedly increased the percentage of resistant aphids in the population, 4 days prior to the chlorpyrifosmalathion mixture treatment. Buschman and DePew (1990) showed that sorghum sprayed with chlorpyrifos and parathion for greenbug control had significantly higher densities of Oligonychus pratensis (Banks grass mite) than untreated fields, but the cause of the mite outbreaks was not determined. Other insecticides have been developed in the past 10 years, such as the soil-applied systemic chloro-nicotinyl, imidacloprid, and the newly developed nicotinamide, Ncyanomethyl-4-trifluromethyl nicotinamide (FMC, 2002). These insecticides, especially when formulated as seed treatments, have gained wide acceptance, and in some areas of the Great Plains have replaced chlorpyrifos as the insecticide of choice (C.D. Patrick, personal communication).

Biological control Using predators and parasites to control greenbug began long before the aphid became a key sorghum pest. Fenton and Dahms (1951) attempted inundative releases of Hippodamia convergens (convergent ladybird) in wheat, but concluded that these were ineffective. Attempts have been made to import and establish predaceous coccinellids and parasitic Hymenoptera to control greenbugs in sorghum and wheat (Jackson et al., 1971; Cartwright et al., 1977; Gilstrap et al., 1984; Michels and Bateman, 1986). However, little success has been achieved in the past 30 years.

Conservation biological control, the use of tactics and approaches that involve the manipulation of the environment of natural enemies so as to enhance their survival, and/or physiological and behavioural performance (Barbosa, 1998), is the current practice of choice. In the southern Great Plains area of the USA, Kring et al. (1985) and Rice and Wilde (1988) demonstrated conclusively that indigenous coccinellids (H. convergens, Hippodamia sinuata, Coleomegilla maculata lengi, and Scymnus spp.) were key to suppressing greenbugs on sorghum early in the season. The impact of the most abundant parasitoid, Lysiphlebus testaceipes, in these experiments was sporadic, typically occurring late in the season. In Nebraska, Fernandes et al. (1998) concluded that L. testaceipes could control greenbugs effectively in an inundative biocontrol programme at a release rate of 24,00036,000 wasps/ha, and suggested that planting alternating strips of greenbug-resistant sorghum hybrids with greenbug-susceptible hybrids as banker plants for L. testaceipes may be an economically feasible way to produce the desired numbers of wasps. There is strong evidence that, at least in the US Great Plains region, greenbug biocontrol by predaceous coccinellids is enhanced when R. maidis is present early in the growing season. Kring and Gilstrap (1986) noted that corn leaf aphids helped maintain Hippodamia spp. in sorghum, and Michels and Behle (1992) found in a 3-year experiment that greenbugs did not reach economic thresholds in 2 years when maize leaf were present early in the season, but severely damaged sorghum in the 1 year that R. maidis populations failed to develop. Field data (G.J. Michels, unpublished results) showed that in 31 irrigated and dryland sorghum fields sampled from 19882000, peak greenbug density never reached damaging levels when corn leaf aphid densities reached 100 or more per plant prior to the sorghum reaching boot stage (approximately 15 July) of a given year (Fig. 27.4). Peak coccinellid density was significantly correlated to corn leaf aphid density rather than greenbug density. Thus, the predators impact on mid-season greenbug infestations may be

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14 12
Com leaf aphids Greenbugs Boot Total predators

Ln density per plant

10 8 6 4 2 0

23/6/1988 29/7/1988 7/9/1988

28/6/1989 2/8/1989

30/6/1998 6/8/1998

Fig. 27.4. Seasonal relationship of corn leaf aphids (Rhopalosiphum maidis), greenbugs (Schizaphis graminum), and predaceous coccinellids in irrigated grain sorghum from 19881990 and 19922000, Texas Agricultural Experiment Station, Bushland, Texas.

predetermined by early-season corn leaf aphid populations. As a bonus, corn leaf aphids seldom cause economic damage to sorghum, even when present in large numbers (G.J. Michels, unpublished results). Wilde and Ohiagu (1976) concluded that chemical control of corn leaf aphid in sorghum did not increase yield and, in light of its potential impact on biological control, was usually unwarranted.

Host-plant resistance Development of greenbug-resistant sorghum hybrids began shortly after the advent of biotype C (Hackerott et al., 1969; Wood, 1971; Teetes and Johnson, 1974). Commercial sorghum hybrids became available in 1976 (Morgan et al., 1980). However, greenbug resistance in sorghum, as in wheat, has been ephemeral due to the development of host races or biotypes (Puterka and Peters, 1995). The ability of greenbugs to overcome previously resistant sorghum hybrids has resulted in the designation of greenbug

biotypes I (Harvey et al., 1991; Bowling et al., 1994) and K (Harvey et al., 1997), although previous biotypes, such as biotype E, had also overcome biotype C-resistant sorghum hybrids. Porter et al. (1997) and Anstead et al. (2003) provide reviews of the greenbug biotype concept in wheat and sorghum. Both papers concluded that biotypes were not engendered by resistant hybrids, but rather that resistant hybrids selected for greenbug genotypes that already existed. Regardless of the ability of greenbugs to overcome previously resistant sorghum genotypes, research continues to develop new sources of resistance (Wilde and Tuinstra, 2000). Antibiosis, antixenosis, and tolerance mechanisms have all been identified in sorghum hybrids either singly or in combination (see Bowling and Wilde, 1996 for a review). Harvey et al. (1997) suggested that breeding efforts should concentrate on multigenic greenbug resistance since there was evidence that such hybrids could maintain their resistance to an array of greenbug genotypes. Harvey et al. (1994) and Thindwa and Teetes (1994) noted that temperature played

5/7/2000 10/8/2000

13/6/1990 18/7/1990 29/8/1990

20/7/1992 26/8/1992

23/7/1993 27/8/1993

22/7/1994 24/8/1994

19/7/1995 23/8/1995

11/7/1996 13/8/1996

21/7/1997 25/8/1997

27/7/1999

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a role in resistance expression. Higher temperatures resulted in delayed development, reduced fecundity, and shorter overall lifespan of greenbugs on resistant sorghum hybrids than on susceptible hybrids or resistant hybrids at lower temperatures. Unfortunately, Rice and Wilde (1989) noted a negative interaction between resistant sorghum hybrids and predaceous coccinellids preying on greenbugs. They observed that H. convergens larval-pupal survival was reduced and eclosion to pupation time was increased when the beetles fed on aphids reared on antibiotic plants. They concluded that the widely accepted concept that hostplant resistance and biological control are compatible was probably too broad a generalization, and that understanding the effects of resistant sorghum hybrids at the third trophic level was essential.

Cultural control Like most crop-feeding insects, greenbugs respond to plant condition and agronomic practices. Schweissing and Wilde (1979) and Archer et al. (1982) found that increasing N fertilizer improved sorghum for the greenbug. Greenbug densities were higher on plants that received N than on those that did not (Schweissing and Wilde, 1979). Archer et al. (1982) noted that greenbugs responded positively to increasing N (0, 45, 90,135, 180, and 225 kg N/ha), and that plant damage

remained essentially the same, even though more plant biomass was produced. The general conclusions were that applying a higher than normal rate of N to sorghum in an attempt to raise crop tolerance to infestation was not a positive greenbug management tool. Harvey and Thompson (1988) reported that greenbug densities were consistently lower on sorghum grown at high plant density than on plants grown at low density. Burton et al. (1987) demonstrated that increasing plant residues on the soil surface by reduced tillage resulted in decreased greenbug density and decreased plant damage. A dense plant canopy that obscured furrows also reduced greenbug infestation. Kindler and Staples (1981) concluded that the greenbug economic injury level was lower in water-stressed than in well-watered sorghum. Michels and Undersander (1986) demonstrated that water stress negatively influenced greenbug reproduction in sorghum when water potential on stressed plants fell below 0.3 MPa. In a 3-year study comparing plant populations and irrigation regimes, Michels et al. (2002) found that peak greenbug densities were highest in well-watered fields with low plant populations, and significantly lower in well-watered fields with high plant populations. The results also indicated that low plant populations coupled with heavy irrigation created more of a greenbug problem than higher plant populations and moderate irrigation amounts (Fig. 27.5).

GREENBUGS/PLANT

150 300
60
80000 160000 320000 240000 HA PLANTS / 400000

450

Fig. 27.5. Graphic representation of greenbug densities associated with varying plant population and total water applied through the season in grain sorghum at Bushland, Texas, 19982000.

10 20 ) 30 (CM 40 ER 50 AT LW TA TO

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Putting the pieces together It is evident from the above discussion that the components for greenbug IPM are available, if somewhat scattered. It is also apparent that the majority of the research into greenbug IPM components has centred in the Great Plains states of North America. A lack of research from other countries may be due to the fact that more important pests have superseded the greenbug (Abate et al., 2000). In the USA, several researchers have published papers that have incorporated at least two IPM components. Starks et al. (1972) conducted a greenhouse experiment that addressed the interaction between resistant sorghum and parasitism by L. testaceipes. They concluded that host-plant resistance and parasitism were complementary factors in reducing greenbug numbers. These results were supported by Dogramaci (1998), who concluded that L. testaceipes parasitism and the use of currently available resistant hybrids were compatible and complimentary control strategies for biotype I greenbugs. In field experiments comparing greenbug densities on resistant and susceptible sorghum hybrids grown at differing plant densities over 2 years, Harvey and Thompson (1988) found that growing resistant hybrids in thick stands reduced greenbug densities significantly over any other combination of hybrid and stand. Susceptible hybrids grown in thick stands yielded similarly to resistant hybrids in thin stands; therefore, the authors concluded that any practice that reduced plant stand would intensify greenbug problems. Burton et al. (1990) extrapolated their previous work (Burton et al., 1987) to compare tillage practices and the inclusion of resistant sorghum hybrids. In conventional tillage plots, greenbug density was significantly lower (50%) on the resistant hybrid. When the resistant hybrid was grown in no tillage plots, greenbug density was significantly lower than on the resistant hybrid grown in conventional tillage plots. A survey and economic analysis of the results by Dharmaratne et al. (1986) encompassed 5 years of sorghum production by farmers in the Texas Blacklands focusing on

the use of resistant hybrids and insecticides to control greenbugs. Texas Blacklands farmers preferred greenbug-resistant sorghums and reported that, without insecticides, there was a net return of approximately US$165/ha using susceptible hybrids without insecticides as against US$200/ha using resistant hybrids. When an average of 1.3 insecticide applications were made, the net returns for susceptible and resistant hybrids were almost the same, US$199 and US$203/ha, respectively. The results demonstrated that the only significant difference in net return occurred between the use of greenbug-susceptible hybrids and all other management strategies. Therefore, farmers would be better off by planting resistant sorghum hybrids and foregoing insecticide treatments. The study also noted that decreasing insecticide use could forestall insecticide resistance, thus preventing increased insecticide use in the future, which would erode net profits. Beneficial effects to the environment through decreased insecticide use were noted, but no monetary value was associated with the effects.

The Current Model and the Future

The best current IPM strategy to control greenbugs on sorghum would be to plant dense stands of resistant hybrids. The plants should receive only the N fertilizer normally used for sorghum production, and water stress should be minimized. Rhopalosiphum maidis, predators, and parasitoids should be monitored to determine if sufficient naturally occurring biological control will be present midway through the growing season. Chemical control should not be prophylactic and should not be employed for corn leaf aphid control. If chemical control does become necessary (determined by localarea economic injury levels), a single application of chlorpyrifos plus malathion could be made to prevent yield loss. Greenbug IPM in sorghum will change over time. Resistant hybrids seem to be the base strategy. However, the history of greenbug biotypes almost guarantees that presently unknown greenbug genotypes will be

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selected for by the currently available resistant hybrids, and breeding new hybrids will continue. New hybrids will undoubtedly have differing agronomic characteristics that may change planting densities and fertilization rates. We would not expect changes in hybrids to influence the benefits accrued through no-till operations. New hybrids may influence biological control, and monitoring the impact new hybrids may have on the third trophic level is important. In our opinion, it would be of great value if this monitoring were incorporated routinely in developing new hybrids rather than an aspect to be studied only after hybrids are released. Chemical control will change dramatically. Organophosphate insecticides will disappear over time and replacements utilizing new chemistries will appear. Given continual environmental concerns, it is likely that these new insecticides will be more specific, less toxic to non-target organisms, and have little residual activity. Therefore, when chemical control is selected, timing will be very important.

utilized. Research scientists must engage extension and consultant personnel to convey information to producers, and actively seek feedback regarding success, failure, and acceptance of recommended strategies.

Executive Summary
In the Great Plains region of the USA, there is currently a fairly complete IPM programme for S. graminum on sorghum for farmers who wish to use it, though adaptation by the aphid to pesticides and resistant varieties is bound to lead to future modifications. Economic thresholds have been established so that prophylactic use of pesticides can be avoided, though the organophosphates for which such an approach is eminently possible are likely to be phased out. Farmers are already using the newer neonicotinoids, and here the simplicity of in-furrow and seed treatment is likely to increase the use of prophylaxis. Indigenous natural enemies can be supported by providing reserve prey in the form of R. maidis, which itself is not a pest problem. Host-plant resistance is the foundation of the IPM programme for greenbug on sorghum, especially when grown at high plant density and irrigation. Cultural control therefore focuses on planting density and water management. There is no use of semiochemicals. These approaches exploit interactions between them, particularly between plant resistance and cultural measures. Unfortunately, it appears that aphid-resistant sorghums may have negative effects on coccinellids by reducing their survival and slowing their development.

Conclusions
Although there are limited case studies addressing greenbug IPM for sorghum, there is a rich literature describing numerous strategies that form a productive whole. There is a need for new research in the form of case studies where single and multiple tactic IPM strategies are compared. When such research is attempted, it is crucial to include economic evaluations and farmer input in the planning stage. With the explosive growth of the Internet, and the resultant availability of information, transfer of research results is easier than in the past, and should be

References
Abate, T., van Huis, A. and Ampofo, J.K.O. (2000) Pest management strategies in traditional agriculture: an African perspective. Annual Review of Entomology 45, 631659. Anstead, J., Burd, J.D. and Shufran, K.A. (2003) Over-summering and biotypic diversity of Schizaphis graminum (Homoptera: Aphididae) populations on non-cultivated grass hosts. Environmental Entomology 32, 662667.

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Archer, T.L., Onken, A.B., Matheson, R.L. and Bynum, E.D., Jr. (1982) Nitrogen fertilizer influence on greenbug (Homoptera: Aphididae) dynamics and damage to sorghum. Journal of Economic Entomology 75, 695698. Archer, T.L., Segarra, E. and Bynum, E.D., Jr. (1999) Greenbug resistance management on sorghum with insecticide mixtures: a biological and economic analysis. Journal of Economic Entomology 92, 794803. Barbosa, P. (1998) Conservation Biological Control. Academic Press, San Diego, 396 pp. Barbulescu, A. (1964) Research on bionomics and control of aphids injurious to sorghum. Problems in Agriculture 16, 2937. Bowling, R. and Wilde, G. (1996) Mechanisms of resistance in three sorghum cultivars resistant to greenbug (Homoptera: Aphididae) biotype I. Journal of Economic Entomology 89, 558561. Bowling, R., Wilde, G., Harvey, T., Sloderbeck, P., Bell, K.O., Morrison, W.P. and Brooks, H.L. (1994) Occurrence of greenbug (Homoptera: Aphididae) biotypes E and I in Kansas, Texas, Nebraska, Colorado, and Oklahoma. Journal of Economic Entomology 87, 16961700. Burton, R.L., Jones, O.R., Burd, J.D., Wicks, G.A. and Krenzer, E.G., Jr. (1987) Damage by greenbug (Homoptera: Aphididae) to grain sorghum as affected by tillage, surface residues, and canopy. Journal of Economic Entomology 80, 792798. Burton, R.L., Burd, J.D., Jones, O.R. and Wicks, G.A. (1990) Crop production strategies for managing greenbug (Homoptera: Aphididae) in sorghum. Journal of Economic Entomology 83, 24762479. Buschman, L.L. and DePew, L.J. (1990) Outbreaks of Banks grass mite (Acari: Tetranychidae) in grain sorghum following insecticide applications. Journal of Economic Entomology 83, 15701574. Cartwright, B.O., Eikenbary, R.D., Johnson, J.W., Farris, T.N. and Morrison, R.D. (1977) Field release and dispersal of Menochilus sexmaculatus, and imported predator of the greenbug, Schizaphis graminum. Environmental Entomology 6, 699704. Cate, J.R., Bottrell, D.G. and Teetes, G.L. (1973a) Management of the greenbug on grain sorghum. 1. Testing foliar treatments of insecticides against greenbugs and corn leaf aphids. Journal of Economic Entomology 66, 945951. Cate, J.R., Bottrell, D.G. and Teetes, G.L. (1973b) Management of the greenbug on grain sorghum. 2. Testing seed and soil treatments for greenbug and corn leaf aphid control. Journal of Economic Entomology 66, 953959. DePew, L.J. (1971) Evaluation of foliar and soil treatments for greenbug control on sorghum. Journal of Economic Entomology 64, 169172. DePew, L.J. (1974) Controlling greenbugs in grain sorghum with foliar and soil insecticides. Journal of Economic Entomology 67, 553555. Dharmaratne, G.S., Lacewell, R.D., Stoll, J.R. and Teetes, G.L. (1986) Economic impact of greenbug resistant grain sorghum varieties: Texas Blacklands. Texas Agricultural Experiment Station Miscellaneous Publication MP-1585, 43 pp. Dibb, D.W. (1983) Agronomic systems to feed the next generation. Crops and Soils Magazine November 1983, 56. Dogramaci, M. (1998) Interaction of Lysiphlebus testaceipes (Cresson) and greenbug (Schizaphis graminum (Rondani)) on resistant and susceptible sorghums. MSc thesis, University of Nebraska, Lincoln, USA. FAO (1979) Production Yearbook, Volume 33. FAO, Rome, 309 pp. FAO (2001) FAOSTAT Database Collections (www.apps.fao.org/). Fenton, F.A. and Dahms, R.G. (1951) Attempts at controlling the greenbug by the importation and release of lady beetles in Oklahoma. Proceedings of the Oklahoma Academy of Science 32, 12. Fernandes, O.A., Wright, R.J. and Mayo, Z.B. (1998) Parasitism of greenbug (Homoptera: Aphididae) by Lysiphlebus testaceipes (Hymenoptera: Braconidae) in grain sorghum: implications for augmentative biological control. Journal of Economic Entomology 91, 13151319. FMC (2002) Flonicamid. FMC Technical Bulletin. FMC, Philadelphia, 6 pp. Gilstrap, F.E., Brooks, G.W. and Kring, T.J. (1984) Status of greenbug biological control in Texas sorghum. Texas Agricultural Experiment Station Miscellaneous Publication MP-1585, 6 pp. Hackerott, H.L., Harvey, T.L. and Ross, W.M. (1969) Greenbug resistance in sorghums. Crop Science 9, 656658. Harvey, T.L. and Hackerott, H.L. (1969) Recognition of a greenbug biotype injurious to sorghum. Journal of Economic Entomology 62, 776779. Harvey, T.L. and Thompson, C.A. (1988) Effects of sorghum density and resistance on infestations of greenbug, Schizaphis graminum (Homoptera: Aphididae). Journal of the Kansas Entomological Society 61, 6871. Harvey, T.L., Kofoid, K.D., Martin, T.J. and Sloderbeck, P.E. (1991) A new greenbug virulent to E biotype resistant sorghum. Crop Science 31, 16891691.

636

G.J. Michels, Jr. and J.D. Burd

Harvey, T.L., Wilde, G.E., Kofoid, K.D. and Bramel-Cox, P.J. (1994) Temperature effects on resistance to greenbug (Homoptera: Aphididae) biotype I in sorghum. Journal of Economic Entomology 87, 500503. Harvey, T.L., Wilde, G.E. and Kofoid, K.D. (1997) Designation of a new greenbug, biotype K, injurious to resistant sorghum. Crop Science 37, 989991. Hunter, S.J. (1909) The greenbug and its enemies. Bulletin of the University of Kansas, No. 9, 213 pp. Jackson, H.B., Rogers, C.E. and Eikenbary, R.D. (1971) Colonization and release of Aphelinus asychis, an imported parasite of the greenbug. Journal of Economic Entomology 64, 14351438. Kindler, S.D. and Staples, R. (1981) Schizaphis graminum: effect on grain sorghum exposed to severe drought stress. Environmental Entomology 10, 247248. Kring, T.J. and Gilstrap, F.E. (1986) Beneficial role of corn leaf aphid, Rhopalosiphum maidis (Fitch) (Homoptera: Aphididae), in maintaining Hippodamia spp. (Coleoptera: Coccinellidae) in grain sorghum. Journal of Crop Protection 5, 125128. Kring, T.J., Gilstrap, F.E. and Michels, G.J., Jr. (1985) Role of indigenous coccinellids in regulating greenbugs (Homoptera: Aphididae) on Texas grain sorghum. Journal of Economic Entomology 78, 269273. Matthee, J.J. (1962) Guard against aphids on kafficorn. Farming in South Africa 37, 2729. Michels, G.J., Jr. and Bateman, A.C. (1986) Larval biology of two imported predators of the greenbug, Hippodamia variegata Goetz and Adalia flavomaculata DeGeer under constant temperatures. Southwestern Entomologist 11, 2330. Michels, G.J., Jr. and Behle, R.W. (1992) Evaluation of sampling methods for lady beetles (Coleoptera: Coccinellidae) in grain sorghum. Journal of Economic Entomology 85, 22512257. Michels, G.J., Jr. and Undersander, D.J. (1986) Temporal and spatial distribution of the greenbug (Homoptera: Aphididae) on sorghum in relation to water stress. Journal of Economic Entomology 79, 12211225. Michels, G.J., Jr., Rush, C.M., Piccinni, G., Fritts, D.A. and Jones, D. (2002) Effect of irrigation regimes and plant population on greenbug (Homoptera: Aphididae) density in grain sorghum. Southwestern Entomologist 27, 135147. Morgan, J., Wilde, G. and Johnson, D. (1980) Greenbug resistance in commercial sorghum hybrids in the seedling stage. Journal of Economic Entomology 73, 510514. Niemczyk, H.D. and Moser, J.R. (1982) Greenbug occurrence and control on turfgrasses in Ohio. In: Niemczyk, H.D. and Joyner, B.G. (eds) Advances in Turfgrass Entomology. Hammen Graphics, Piqua, Ohio, pp. 105111. Peters, D.C., Wood, E.A., Jr. and Starks, K.J. (1975) Insecticide resistance in selections of the greenbug. Journal of Economic Entomology 68, 339340. Porter, D.R., Burd, J.D., Shufran, K.A., Webster, J.A. and Teetes, G.L. (1997) Greenbug (Homoptera: Aphididae) biotypes: selected by resistant cultivars or preadapted opportunists? Journal of Economic Entomology 90, 10551065. Puterka, G.J. and Peters, D.C. (1995) Genetics of greenbug (Homoptera: Aphididae) virulence to resistance in sorghum. Journal of Economic Entomology 88, 421429. Rice, M.E. and Wilde, G.E. (1988) Experimental evaluation of predators and parasitoids in suppressing greenbugs (Homoptera: Aphididae) in sorghum and wheat. Environmental Entomology 17, 836841. Rice, M.E. and Wilde, G.E. (1989) Antibiosis effect of sorghum on the convergent lady beetle (Coleoptera: Coccinellidae), a third-trophic level predator of the greenbug (Homoptera: Aphididae). Journal of Economic Entomology 82, 570573. Rider, S.D., Dobesh, S.M. and Wilde, G.E. (1998) Insecticide resistance in a novel laboratory-produced greenbug (Homoptera: Aphididae) clone. Journal of Economic Entomology 91, 3033. Schweissing, F.C. and Wilde, G. (1979) Temperature and plant nutrient effects on resistance of seedling sorghum to the greenbug. Journal of Economic Entomology 72, 2023. Shufran, R.A., Wilde, G.E. and Sloderbeck, P.E. (1996) Descriptions of three isozyme polymorphisms associated with insecticide resistance in greenbug (Homoptera: Aphididae) populations. Journal of Economic Entomology 89, 4650. Shufran, R.A., Wilde, G.E. and Sloderbeck, P.E. (1997a) Response of three greenbug (Homoptera: Aphididae) strains to five organophosphorus and two carbamate insecticides. Journal of Economic Entomology 90, 283286. Shufran, R.A., Wilde, G.E. and Sloderbeck, P.E. (1997b) Life history of insecticide resistant and susceptible greenbug (Homoptera: Aphididae) strains. Journal of Economic Entomology 90, 15771583. Sloderbeck, P.E., Chowdhury, M.A., DePew, L.J. and Buschman, L.L. (1991) Greenbug (Homoptera: Aphididae) resistance to parathion and chlorpyrifos-methyl. Journal of the Kansas Entomological Society 64, 14.

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Starks, K.J., Muniappan, R. and Eikenbary, R.D. (1972) Interaction between plant resistance and parasitism against the greenbug on barley and sorghum. Annals of the Entomological Society of America 65, 650655. Stone, B.S., Shufran, R.A. and Wilde, G.E. (2000) Life history study of multiple clones of insecticide resistant and susceptible greenbug Schizaphis graminum (Homoptera: Aphididae). Journal of Economic Entomology 93, 971974. Teetes, G.L. and Johnson, J.W. (1974) Assessment of damage by the greenbug in grain sorghum hybrids of different maturities. Journal of Economic Entomology 67, 514516. Teetes, G.L., Schaefer, C.A., Gipson, J.R., McIntyre, R.C. and Latham, E.E. (1975) Greenbug resistance to organophosphorus insecticides on the Texas High Plains. Journal of Economic Entomology 68, 214216. Thindwa, H.P. and Teetes, G.L. (1994) Effect of temperature and photoperiod on sorghum resistance to biotype C and E greenbug (Homoptera: Aphididae). Journal of Economic Entomology 87, 13661372. USDA (1968) Greenbug (Schizaphis graminum). Plant Pest Control Division Cooperative Insect Report, No. 18, 781 pp. USDA (1981) Components for management of field corn, and grain sorghum insects and mites in the United States. USDA-ARS ARM-S-18. USDA-ARS, New Orleans, 18 pp. Webster, J.A., Amosson, S., Brooks, L., Hein, G., Johnson, G., Legg, D., Massey, B., Morrison, P., Peairs, F. and Weiss, M. (1995) Economic impact of the greenbug in the western United States: 19921993. Great Plains Council Publication, No. 155, 16 pp. Wilde, G.E. and Ohiagu, C. (1976) Relation of corn leaf aphid to sorghum yield. Journal of Economic Entomology 69, 195- 197. Wilde, G.E. and Tuinstra, M.R. (2000) Greenbug (Homoptera: Aphididae) resistance in sorghum: characterization of KS 97. Journal of Agricultural and Urban Entomology 17, 1519. Wood, E.A. (1971) Designation and reaction of three biotypes of the greenbug cultures on resistant and susceptible species of sorghum. Journal of Economic Entomology 64, 183185. Young, W.R. and Teetes, G.L. (1977) Sorghum entomology. Annual Review of Entomology 22, 193218.