Effect of cropping systems on cereal stemborers in the cool‐wet and semi‐arid ecozones of the Amhara region of Ethiopia

1 Field experiments were conducted on maize and sorghum at three locations in the Amhara state of Ethiopia to determine the effects of mixed cropping on stemborer infestation, borer natural enemies and grain yields. In the cool‐wet ecozone of western Amhara, sole maize was compared with maize intercropped with faba bean, mustard, potatoes and cowpea. In the semi‐arid ecozone of eastern Amhara, the trial was conducted on both maize and sorghum with the companion crops haricot bean, sesame, cowpea and sweet potatoes. 2 The results showed that the predominant borer species in western and eastern Amhara were, respectively, Busseola fusca and Chilo partellus. In Addis Zemen, western Amhara, maize intercropped with mustard and potatoes had significantly lower pest numbers and percent tunnelling than other intercrops and the maize monocrop during the vegetative stage. In eastern Amhara, the cropping system did not significantly affect pest densities but damage to stem, ear or heads tended to be greatest when cereals were intercropped with sweet potatoes. 3 Parasitism of C. partellus by the braconid Cotesia flavipes was greater on maize than sorghum, and on maize it was greater with sweet potatoes than in other intercrops or sole maize. Cocoon mass number per plant did not vary significantly between treatments. 4 There were significant differences between treatments in yields of both sorghum and maize (per plant and per unit area) with the lowest yields observed when they were intercropped with a tuber crop. 5 The results suggest that simultaneous planting of the crop species selected has little advantage over monocropped maize.     

Distribution and relative importance of cereal stem borers and their natural enemies in the semi-arid and cool-wet ecozones of the Amhara State of Ethiopia

The distribution and relative importance of lepidopteran and coleopteran stem borers and their natural enemies on maize and sorghum were studied in cereal growing zones of the Amhara State of Ethiopia from 2003 to 2004. Sorghum is the major crop in semi-arid eastern and maize in the cool-wet western zones of the Amhara state. Four administrative zones, 10 districts and 88 localities in the semi-arid ecozone (SAE) and four zones, 19 districts and 71 localities in the cool-wet ecozone (CWE) were chosen for the study. In SAE, the species composition was 91% Chilo partellus (Swinhoe) (Lepidoptera: Crambidae), 8% Busseola fusca (Fuller) (Lepidoptera: Noctuidae) and 1% Sesamia calamistis Hampson (Lepidoptera: Noctuidae). In the CWE, maize and sorghum are grown in different ecozones and thus B. fusca was the dominant species on sorghum, whereas 61% B. fusca and 39% S. calamistis were recorded on maize. Borer density generally increased with crop growth stage. C. partellus parasitism by C. flavipes Cameron (Hymenoptera: Braconidae), which occurred only in SAE, varied among districts ranging from 5% to 39%. In the CWE, unidentified nematodes parasitized medium-sized B. fusca larvae during the wet months. Population of native parasitoids was very low. The coleopteran borer, Rhynchaenus niger (Horn) (Coleoptera: Rhynchophoridae), attacked sorghum plants in both regions. Sorghum yields were negatively related to plant damage variables and positively to larval parasitism and plant growth variables. On maize, plant damage was too low to affect yields. Taylor’s power law indicated aggregated distribution for C. partellus and B. fusca larvae and pupae combined.     

Yield loss due to the stemborer Chilo partellus (Swinhoe) (Lepidoptera: Crambidae) at different nitrogen application rates to maize

Field trials were conducted at Kibaha and Morogoro in eastern Tanzania during two seasons to evaluate the effect of nitrogen fertilization (0, 50, 75, 100 kg [N]/ha) on pest abundance, plant damage and yield loss of maize due to stemborers. In general, ear and grain weights increased linearly with nitrogen level. In the infested plot, grain weight increased 2.5 and 1.8 fold from 0 to 100 kg [N]/ha in the short and long rainy season, respectively, at Kibaha, and 1.4 and 1.6 times at Morogoro. Yield loss decreased with an increase in nitrogen application and the effect was stronger under high than low borer infestation levels. The results show the beneficial effect of nitrogen on the plant’s ability to compensate for borer damage. Analysis of economic benefits of applying fertilizer and insecticide treatment indicated that using insecticides is not profitable under high-pest-low-soil fertility conditions.     

The effect of grassy field margins and fertilizer on soil water,plant nutrient levels,stem borer attacks and yield of maize in the humid forest zone of Cameroon

Two field experiments, planted in a split plot design, were conducted during 2002 in the forest zone of Cameroon, to investigate the effect of border rows with Pennisetum purpureum (Poaceae) or with Panicum maximum (Poaceae) on soil water, plant nitrogen (N), phosphorus (P) and potassium (K), borer infestations, parasitism and maize yield. The grassy boundaries were the main plots and fertilizer treatment the sub plots. Soil humidity was significantly higher under the grass borders than in maize plots. Nitrogen uptake by maize tended to be highest in plots surrounded by P. purpureum but the differences were significant during the second season only. Busseola fusca (Fuller) (Lepidoptera: Noctuidae) was the predominant borer species followed by Eldana saccharina (Walker) (Lepidoptera: Pyralidae). The predominant parasitoid species was the scelionid egg parasitoid Telenomus busseolae Gahan (Hymenoptera: Scelionidae). During both seasons, plant nitrogen, B. fusca infestation, plant damage and yield were significantly higher in maize sub-plots that received fertilizer The interaction between the grassy margin effect and the fertilization was significant only for B. fusca infestations, during the second season with maize + P. purpureum having a significantly lower number of borers in the fertilized than the unfertilized sub-plots. The grassy borders had no effect on B. fusca egg parasitism variables except in the first season, when maize with P. purpureum had a significantly higher percentage of egg batches parasitized. During both seasons, there were some significant differences in yield variables between main plots but the trends were not clear. Multiple regression showed that B. fusca infestation, plant damage, egg parasitism, plant N, P and K affected yield, with plant nutrients explaining most of the variability. The implication of the findings for the feasibilty of this habitat management technology to farmers in southern Cameroon is discussed.     

The role of intercropping different cereal species in controlling lepidopteran stemborers on maize in Kenya

Abstract:  The effects of mixed cropping systems containing maize, sorghum, millet and beans on infestations of cereals by lepidopteran stemborers and on associated parasitoids, as well as on yields and land equivalent ratios (LER) were assessed during four consecutive rainy seasons at two sites in the semi-arid eastern region of Kenya. Systems containing the non-host bean were more efficient in reducing pest densities than those with millet or sorghum only. Higher parasitism in diversified systems compared to monocrops was due to density-dependent effects rather than superior suitability of such systems to parasitoids. The maize–bean system, which had the highest proportion of bean plants, had LERs >1.65 while most other systems had LERs <1. It is concluded that mixed cropping with several cereal species has little advantages in terms of yield loss abatement due to stemborers and land use efficiency. However, including the drought-tolerant crops such as sorghum and millet in the system stabilizes food security in drought-prone areas such as eastern Kenya.     

Stem borer infestation of cereal crops in Jere,the Sudan-Sahelian savanna region of Nigeria

Field data collected during the rainy season of two years, 2010 and 2011, were used to determine the per cent plant infestation and stem borer abundance on cultivated cereal crops grown by farmers' in Jere or the Sudan-Sahelian savanna ecological region of Nigeria. Stem borers were recovered using destructive sampling. Mean total per cent plant infestation and stem borer abundance per farmers' field were significantly higher on millet (40% and 25 individuals, respectively) and sorghum (30% and 21 individuals, respectively) than on maize (19% and 13 individuals, respectively). Of the five stem borer species found in this study, Coniesta ignefusalis (Lepidoptera: Pyralidae) (3.5)/Chilo sp. nr. aleniellus (Lepidoptera: Crambidae) (2), Busseola fusca (Lepidoptera: Noctuidae) (3.1)/Sesamia calamistis (Lepidoptera: Noctuidae) (2.4) and S. calamistis (2.9), with significantly higher number of individuals per plant, tended to be more important pests on millet, sorghum and maize crops, respectively. Although, mean total per cent plant infestation and abundance of stem borers in this study were generally moderate, further studies on the effects of different types of cereals intercropping (locally practiced) on stem borer infestation and abundance should ascertain the true importance of these pest species in the Sudan-Sahelian savanna ecological region of Nigeria.     

Forest bolsters bird abundance,pest control and coffee yield

Efforts to maximise crop yields are fuelling agricultural intensification, exacerbating the biodiversity crisis. Low‐intensity agricultural practices, however, may not sacrifice yields if they support biodiversity‐driven ecosystem services. We quantified the value native predators provide to farmers by consuming coffee's most damaging insect pest, the coffee berry borer beetle (Hypothenemus hampei). Our experiments in Costa Rica showed birds reduced infestation by ~ 50%, bats played a marginal role, and farmland forest cover increased pest removal. We identified borer‐consuming bird species by assaying faeces for borer DNA and found higher borer‐predator abundances on more forested plantations. Our coarse estimate is that forest patches doubled pest control over 230 km² by providing habitat for ~ 55 000 borer‐consuming birds. These pest‐control services prevented US$75–US$310 ha‐year⁻¹ in damage, a benefit per plantation on par with the average annual income of a Costa Rican citizen. Retaining forest and accounting for pest control demonstrates a win–win for biodiversity and coffee farmers.     

Mechanisms and genetic diversity for host plant resistance to spotted stem borer,Chilo partellus in sorghum,Sorghum bicolor

Levels of resistance to spotted stem borer, Chilo partellus, in the germplasm are low to moderate and therefore we evaluated 25 sorghum genotypes for resistance to stem borer to identify lines with diverse mechanisms of resistance to this insect. Leaf glossiness was significantly and negatively associated with low deadheart formation. Dwarf genotypes with fewer nodes showed more deadheart formation. Antixenosis and/or antibiosis for leaf feeding, and reduced deadheart formation, tunnelling, and number of exit holes resulted in lower losses owing to C. partellus damage. Axillary tillers compensated for loss in grain yield owing to borer infestation as a result of deadheart formation, but their synchrony for maturity with the main plant is quite important. Path coefficient analysis revealed that direct effects of stem tunnelling on loss in grain yield were greater than leaf feeding and deadhearts. However, leaf feeding via stem tunnelling showed maximum indirect effects on loss in grain yield. Estimates for broad‐sense heritability and genetic advance suggested that it is possible to improve for grain yield under stem borer infestation. Heritability estimates for grain yield were usually higher than for stem borer damage parameters. Multi‐trait cluster analysis placed the test genotypes into four and five clusters in rainy and post‐rainy seasons, respectively. The genotypes placed in different groups, and showing resistance to leaf feeding, deadheart formation, stem tunnelling, and/or compensation in grain yield can be used for sorghum improvement.     

Onion fly egg distribution in soil: a new sampling method and the effect of two granular insecticides

Tests were carried out in a screenhouse to determine the potential of a microsporidian pathogen,Nosema marucae (Microspora: Nosematidae), for the control of the spotted stalk borer,Chilo partellus (Lepidoptera: Pyralidae), on sorghum. Foliar application of an aqueous suspension ofN. marucae spores on egg batches pasted on the sorghum plants reduced leaf-damage in plants in the protected plots, and increased the proportion of plants with fully- and partially formed heads compared to unsprayed plots. Further, there was less tunnelling and fewer larvae in the sprayed plots. The pathogen also reduced pest infestation and damage to plants when the plots were infested with newly emerged larvae instead of eggs.N. marucae has a potential to controlC. partellus.     

Disturbance of the rainforest has the potential to enhance egg parasitism of lepidopteran noctuid stemborers in Kisangani,DR Congo

Landscape context influences population dynamics of insects and impacts biological processes within communities. It was expected that anthropogenic disturbances of the rainforest landscape in DR Congo would lead to a decreased level of noctuid stemborer egg parasitism as a consequence of a decoupling between stemborers and their naturally occurring parasitoids through dispersal. To test this hypothesis, noctuid egg batches were collected in maize fields along an anthropogenic disturbance gradient to assess change in the rates of eggs parasitism and maize plant infestation with noctuid egg batches. Our results showed that, in contrast to what was initially expected, egg parasitism increased from less to highly disturbed landscape whereas maize infestation had an inverse tendency. Discovery efficiency and mean egg parasitism were 1.416 and 1.392 times higher, respectively, in the most than in the less disturbed landscape. The numbers of eggs and egg batches per 100 maize plants were 0.55 times and 0.532 times the value in the less disturbed landscape, suggesting a dilution of the stemborer population within a large habitat patch encompassing cultivated fields and the surrounding wild host plants. It was concluded that the presence of suitable host plants enhances noctuid stemborers egg parasitism in adjacent maize fields.     

The effect of nitrogen fertilizer application to maize and sorghum on the bionomics of Chilo partellus (Lepidoptera: Crambidae) and the performance of its larval parasitoid Cotesia flavipes (Hymenoptera: Braconidae)

Laboratory and field trials were conducted to evaluate the effect of plant species (maize, sorghum), plant age (young, middle, old) and four different nitrogen fertilization levels (N0-N3) on the bionomics of the invasive crambid Chilo partellus and the performance of its braconid larval parasitoid Cotesia flavipes. Plant N varied significantly between N0 and N1-N3, but the differences among the latter were not significant. Intrinsic rates of increase and net-reproductive rates of C. partellus followed the same trends: they were lowest with N0 and similar among the other treatments. On maize only, mortality of C. partellus and parasitism by C. flavipes tended to decrease with age of the plant while the percentage of borers reaching adulthood (i.e. pupation) increased. Borer mortality and parasitism was lower and pupation higher on sorghum than on maize. On both host plants, percent dry matter content of frass, which could affect ingress of the parasitoid into the borer tunnel, did not vary with nitrogen level but varied with age of the host plants: on maize, it was highest on young plants and on sorghum on old plants. Tunnels were shorter on young maize and sorghum plants; longer tunnels on older plants indicated compensatory feeding by the larva as a result of lower nutritive value of the food source. Consequently, larval weight was lower on older than younger plants. The level of nitrogen fertilization had no effect on food conversion efficiency of C. partellus. Nitrogen did not affect number of C. flavipes progeny while egg load of progeny increased significantly with nitrogen level, on both plant species. Differences in egg load between sorghum and maize were mostly not significant. It was concluded that on depleted soils only, an increase in nitrogen via mulching, rotation with a leguminous crop or fertilization would increase survival of C. partellus on both maize and sorghum and an increase in acreage of maize and in application of nitrogen fertilizer in an area would also increase the parasitism of C. flavipes.     

Dynamics of infestation and losses of stored maize due to the larger grain borer (Prostephanus truncatus Horn) and maize weevils (Sitophilus zeamais Motschulsky)

The dynamics of infestation and losses of stored maize due to the larger grain borer (Prostephanus truncatus Horn) and maize weevils (Sitophilus zeamais Motschulsky) were investigated in Tanzania. In single species populations, significantly higher maize damage (P ≤ 0.05) was caused by P. truncatus (23.3%) compared to S. zeamais (6.4%). Weight losses caused by P. truncatus were higher (12.7%) than for S. zeamais (6.7%) under similar conditions. A higher proportion of P. truncatus in the initial infestation population led to significantly higher (≤0.05) maize damage compared to equal or higher proportions of S. zeamais. Irrespective of the initial number of S. zeamais, population size increased to levels higher than for P. truncatus. Maize losses in unimproved storage structures were >80% compared to <2.5% in improved structures under natural infestation. Inter-specific competition appeared to influence the dynamics of infestation, damage levels and weight losses. The first two months of storage were critical for damaging insect populations to build up.     

Response of maize stemborers and associated parasitoids to the spread of grasses in the rainforest zone of Kisangani,DR Congo: effect on stemborers biological control

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Associations between host plant concentrations of selected biochemical nutrients and Mexican rice borer,Eoreuma loftini,infestation

The Mexican rice borer, Eoreuma loftini (Dyar) (Lepidoptera: Crambidae), is an economic pest of sugarcane (Saccharum spp. hybrids) and other graminaceous host crops, and it attacks grassy weeds. Oviposition preference has been known to be for plants with leaves that form folds. This study is the first to associate the nutritional quality of crop and forage plant hosts with Mexican rice borer injury. Three experiments were conducted to determine the levels of selected biochemical nutrients, including free amino acids and sugars, in four grass weeds [barnyardgrass, Echinochloa crus‐galli (L.); broadleaf signalgrass, Urochloa platyphylla (Munro ex C. Wright); johnsongrass, Sorghum halepense (L.) Pers.; and Vasey's grass, Paspalum urvillei Steud.], a forage grass [sudangrass, Sorghum bicolor (L.) Moensch spp. drummondi (Nees ex Steud.) de Wet & Harlan], three crop species [maize, Zea mays L.; sorghum, Sorghum bicolor (L.) Moench; and sugarcane, Saccharum spp.], and a sorghum‐sudangrass forage hybrid (all Poaceae). Of 16 free amino acids detected among plants in the first two experiments, only high accumulations of free histidine in sudangrass and maize were associated with increased infestations by Mexican rice borer larvae. In all three experiments, high levels of fructose were associated with heightened Mexican rice borer infestation. Ramifications of these findings on the potential dispersal of this invasive pest in the USA and possible applications of fructose in baits are discussed.     

Effect of nitrogen fertilizer level on infestations of lepidopterous stemborers and yields of maize in Zanzibar

The effect of nitrogen levels of 0, 60, 120, and 250 kg/ha and insecticide treatment (Furadan) on population densities and parasitism of lepidopteran stemborers, and maize yields were studied in Zanzibar during 2004/05. Chilo partellus (Swinhoe) (Lepidoptera: Crambidae) dominated by 3-fold over Sesamia calamistis Hampson (Lepidoptera: Noctuidae) and 42 fold over Chilo orichalcociliellus Strand (Lepidoptera: Crambidae). Stemborer density per plant and parasitism by Cotesia flavipes (Cameron) (Hymenoptera: Braconidae) increased with nitrogen application level. Percentage of bored internodes per plant caused by stemborer decreased with N levels during the short rainy season. Pesticide application reduced densities of all stemborer species during the short rainy season, when infestations were high. Maize yield increased 2 to 8 times with N level, compared to the zero treatment, but the effect was less pronounced in the protected plots.     

Density dependence and temporal plasticity of competitive interactions during utilisation of resources by a community of lepidopteran stemborer species

Interactions within and between species sharing the same resources are characterised by competition or facilitation, and can be influenced by factors such as larval numbers and phenotypic plasticity of the interactions. The effect of larval density on the survival and relative growth rate of the stemborers Busseola fusca (Fuller) and Sesamia calamistis Hampson (both Lepidoptera: Noctuidae), and Chilo partellus (Swinhoe) (Lepidoptera: Crambidae) were studied, as well as the temporal plasticity of their competitive interactions. These stemborers attack maize crops (Zea mays L.) (Poaceae) in sub‐Saharan Africa. Experiments were conducted in the laboratory under controlled conditions at the optimum development temperature (25 °C) for the three species. Surrogate stems filled with artificial diet were intra‐ and interspecifically infested with larvae of each species. The effect of larval density on competition was studied at low (six larvae) and high (12 larvae) levels of infestation, whereas the temporal plasticity of competition was evaluated at 7, 14, 21, or 28 days after infestation. The two experiments involved single‐ and multi‐species infestation treatments. Larval numbers and wet mass in each artificial stem were recorded in each experiment. Survival and relative growth rate of the three species were significantly higher at low‐infestation levels when facing either intra‐ or interspecific competition. The intensity of competition was also temporally plastic among the species and increased as the duration of competition increased. These results are discussed in terms of general infestations of cereal crops by borers.     

Natural enemies of the maize cob borer,Mussidia nigrivenella (Lepidoptera: Pyralidae) in Benin,West Africa

Mussidia nigrivenella Ragonot is a pest of maize cobs in West Africa. It significantly reduces maize yields and grain quality, with quantitative losses of 2-25%at harvest, and up to 10-15% indirect losses due to an increase in storage pest infestation levels. Infestation by M. nigrivenella also significantly increased the susceptibility of maize to Aspergillus flavus infection and subsequent aflatoxin contamination. Surveys conducted in different agro-ecological zones of Benin on cultivated and wild host plants during 1994-1997 revealed one egg parasitoid, three larval parasitoids and one pupal parasitoid attacking M. nigrivenella. Egg parasitism was scarce on all host plants sampled and in all four agro-ecological zones. Parasitism by larval and pupal parasitoids was usually less than 10%, and varied with host plant species. Both larval and pupal parasitoids were rare or absent in cultivated maize fields. The solitary chalcidid pupal parasitoid, Antrocephalus crassipes Masi, was the predominant species, contributing approximately 53% of the observed mortality. Logistic regression analysis indicated that this parasitoid was more prevalent on fruits of Gardenia spp. (Rubiaceae) than on the other host plant species including maize used by M. nigrivenella, and was most abundant between February and September. The differences in parasitoid diversity and parasitism between Benin and other regions suggest that there are opportunities for biological control through introduction of exotic parasitoids or using the 'new association' approach, which uses natural enemies of closely related host species that occupy similar ecological niches to the target pest.     

Labeled nitrogen fertilizer research with urea in the semi-arid tropics

Summary Field studies with bordered microplots were conducted on an Alfisol in the semiarid tropics of India to determine (1) the fate of¹⁵N-labeled urea applied to dryland sorghum in two successive rainy seasons and (2) the effect of method of application on N fertilizer efficiency. Recoveries of¹⁵N-labeled fertilizers by above-ground plant parts ranged from 46.7% to 63.6% in 1981 when the rainfall was above the average and from 54.4% to 66.9% in 1980 when the rainfall was near the average. Small (0.014 g) pellets of urea applied twice as postemergent applications in separate 5 cm deep bands were more effective than single preemergent applications either surface applied or incorporated. Both banding and the split applications contributed to overall fertilizer efficiency. Large (1.0 g) pellets of urea (supergranules) placed at a depth of 5 cm were also superior to the incorporated, small-pellet treatment in 1981. The¹⁵N-balance data for the soil (0–90 cm in depth)-plant system in 1981 showed that the unaccounted-for fertilizer N ranged from 5.1% to 20.6%. An important finding was that high grain yields, in excess of 6,000 kg/ha, with N fertilizer losses of less than 10% could be obtained through fertilizer management during a very wet season. The data from the Alfisol experiments were compared with data from similar Vertisol experiments; N fertilizer losses resulting from incorporated and surface applications were greater for Vertisols than for Alfisols in the wetter year.     

Evidence of the establishment of Cotesia flavipes (Hymenoptera: Braconidae), a parasitoid of cereal stemborers,and its host range expansion in Ethiopia

Three lepidopteran cereal stemborers, Chilo partellus (Swinhoe) (Crambidae), Busseola fusca Fuller, and Sesamia calamistis Hampson (Noctuidae) were collected from maize and sorghum in Ethiopia. The noctuid stemborers are indigenous to Africa while C. partellus is an introduced species from Asia. In 1999, the Asian stemborer parasitoid, Cotesia flavipes Cameron (Braconidae) was found to be widespread in Ethiopia, even though it had never been released in the country. In addition to attacking Chilo partellus, Cotesia flavipes was reared from B. fusca and S. calamistis. The origin of C. flavipes in Ethiopia may have been Somalia where it was released in 1997 near the border with eastern Ethiopia. Percent parasitism of borers by C. flavipes was higher in eastern Ethiopia than other surveyed regions, and parasitism was higher in 2000 than 1999. Parasitism was higher when cereals were intercropped with other plants and when wild grass hosts of stemborers were present.     

Increase in cereal stem borer populations through partial elimination of natural enemies

An insecticide exclusion method was used to evaluate the effect of parasitoids on level of infestation by the stem borers, Busseola fusca (Fuller) and Chilo partellus (Swinhoe), in grain sorghum. In field trials conducted at Brits and at Delmas, South Africa, a selective organophosphate insecticide, dimethoate, was applied twice weekly at each site to three subplots whereas three other identical subplots served as controls. Twelve plants were randomly selected from each subplot at weekly intervals and removed from the field. In the laboratory all plants were dissected to record borer infestation. In order to determine parasitism levels egg batches were kept in Petri dishes and all borer larvae and pupae were kept individually in vials until either parasitoids or moths emerged. At Brits ca. 97% of borers were C. partellus and 3% B. fusca, whereas at Delmas 37.5% were C. partellus and 62.5% B. fusca. The most abundant parasitoids of B. fusca were Cotesia sesamiae (Cameron) and Bracon sesamiae Cameron. The dominant parasitoids of C. partellus at both sites were C. sesamiae, Stenobracon spec., Dentichasmias busseolae Henrich and Pediobius furvus (Gahan). No egg parasitoids were found. At both sites, infestation levels in the sprayed plots were significantly higher than in the untreated plots. On the other hand, parasitism levels of borers in the unsprayed plots were significantly higher than in the treated plots. It was concluded that the higher infestation level of sorghum by stem borers in the sprayed plots was because of partial elimination of parasitoids and possibly other natural enemies by the pesticide.