Sitophilus zeamais (greater grain weevil)
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- Habitat List
- Hosts/Species Affected
- Host Plants and Other Plants Affected
- Growth Stages
- List of Symptoms/Signs
- Biology and Ecology
- Natural enemies
- Notes on Natural Enemies
- Detection and Inspection
- Similarities to Other Species/Conditions
- Prevention and Control
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Sitophilus zeamais Motschulsky
Preferred Common Name
- greater grain weevil
Other Scientific Names
- Calandra chilensis (Philippi & Philippi)
- Calandra oryzae
- Calandra oryzae platensis
- Calandra platensis (Zacher)
- Calandra quadrimacula (Walker)
- Calandra zeamais Motschulsky
- Calandra zemais (Motschulsky)
- Calendra zeamais Motschulsky
- Sitophilus oryzae platensis
- Sitophilus oryzae zeamaiz Motschulsky
International Common Names
- English: billbug, northern corn; maize weevil; weevil, greater rice; weevil, maize
- Spanish: gorgojo del grano; gorgojo del maiz; gorgojo del maíz almacenado; picudo del maíz
- French: calandre du mais
- Portuguese: gorgulho do milho
Local Common Names
- Germany: Kaefer, La-Plata-Mais-; Kaefer, Mais-
- Iran: susske sorrat
- Japan: kokuzo
- Norway: maissnutebille
- Turkey: misir biti
- CALAZM (Sitophilus zeamais)
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Arthropoda
- Subphylum: Uniramia
- Class: Insecta
- Order: Coleoptera
- Family: Dryophthoridae
- Genus: Sitophilus
- Species: Sitophilus zeamais
Notes on Taxonomy and NomenclatureTop of page
First described by Linnaeus in 1798 as Curculio oryza, the first named species of the group was later revised by De Clairville and Scheltenburg in 1798 as Calandra oryzae, which uses the commonest generic synonym for Sitophilus. Many workers subsequently recognized that two distinct forms of the species existed, which were described as the 'large' and 'small' forms. In 1855, Motschulsky recognized the large form as a distinct species, which he named Sitophilus zeamais. Unfortunately, few workers recognized this revision and the name Calandra oryzae continued to be applied to all insects in this complex. Takahashi in 1928 and 1931 complicated matters by raising the small form to specific status as Calandra sasakii. This confused situation continued until 1959, when Floyd and Newsom (1959) revised the complex; this was followed by a further revision by Kuschel (1961). In these revisions it was shown that Linnaeus originally described the smaller species and that Motschulsky's description of the larger species was valid. Both species were therefore placed in the genus Sitophilus with the specific names proposed by Linnaeus and Motschulsky.
Unfortunately, the size difference between S. oryzae and S. zeamais is not consistent, so it is not possible to be sure that references to the large and small forms of Calandra oryzae refer to S. zeamais and S. oryzae, respectively. Therefore the only true and unconfused synonym of S. oryzae is Calandra sasakii; in pre-1960s literature, Calandra oryzae 'small' and 'large' forms could refer to either S. zeamais or S. oryzae, and it is also possible that some references to 'S. oryzae' in the 1960s and early 1970s literature actually relate to S. zeamais misidentified by use of old keys. The genus Sitophilus and its species may be identified using the keys of Gorham (1987) or Haines (1991).
DescriptionTop of page
These developmental stages are all found within tunnels and chambers bored in the grain and are thus not normally seen. The larvae are apodous.
Usually blacker than S. oryzae, with fine microsculpture and is more shiny. Scutellum with lateral elevations further apart than their longitudinal length which is about half as long as the scutellum.
Males with median lobe of aedeagus with two longitudinal grooves dorsally, except in the apical quarter, and is thus sinuous in cross section.
Females with lateral lobes of the Y-shaped sclerite pointed and their separation is greater than for S. oryzae.
S. oryzae and S. zeamais are almost indistinguishable from each other externally; identification is by examination of the genitalia. Both have the characteristic rostrum and elbowed antennae of the family Curculionidae. The antennae have eight segments and are often carried in an extended position when the insect is walking. Both species usually have four pale reddish-brown or orange-brown oval markings on the elytra, but these are often indistinct. (See also S. oryzae.)
Both species can be separated from S. granarius by the presence of wings beneath the elytra (absent in S. granarius) and by having circular, rather than oval, punctures on the prothorax. Molecular characters also separate S. oryzae and S. zeamais and confirm reproductive isolation (Hidayat et al., 1996).
DistributionTop of page
The detailed map plotted on the basis of actual country records gives a falsely restricted distribution. These pests are carried all over the world in grain shipments and can establish themselves wherever there is food and where grain moisture and temperature are favourable. In various locations, one species may be more common than the other. A global survey of resistance to pesticides (Champ and Dyte, 1976) contains detailed location lists for both species.
Distribution TableTop of page
The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.Last updated: 10 Feb 2022
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Central African Republic||Present|
|Congo, Republic of the||Present|
|Federal Republic of Yugoslavia||Present|
|Germany||Present||Introduced||First reported: <1927|
|Sweden||Present||Introduced||First reported: <1995|
|Antigua and Barbuda||Present|
|Trinidad and Tobago||Present|
|United States||Present||Present based on regional distribution.|
|Australia||Present||Present based on regional distribution.|
|-New South Wales||Present|
|Papua New Guinea||Present|
|Brazil||Present||Present based on regional distribution.|
|-Rio de Janeiro||Present|
|-Rio Grande do Sul||Present|
Habitat ListTop of page
Hosts/Species AffectedTop of page
In the case of rice, detailed surveys in Indonesia have shown that S. zeamais is dominant on milled rice, whereas S. oryzae is more common on paddy (rough rice). Laboratory studies have shown that this is a result of their different rates of increase on these two forms of rice (Hussain et al., 1985). It is not yet known whether these relationships with the form of rice hold true throughout the tropics, but imports of milled rice into the UK from many countries are much more frequently infested by S. zeamais than by S. oryzae.
Both species are able to breed on dried cassava and have been reported as frequent pests of this commodity. A few strains of S. oryzae have been found which can develop on grain legumes (Coombs et al., 1977): peas, lentils and green or black gram are the pulses most often attacked by these strains.
Host Plants and Other Plants AffectedTop of page
|Carya illinoinensis (pecan)||Juglandaceae||Unknown|
|Colocasia esculenta (taro)||Araceae||Unknown|
|Glycine max (soyabean)||Fabaceae||Unknown|
|Manihot esculenta (cassava)||Euphorbiaceae||Main|
|Oryza sativa (rice)||Poaceae||Main|
|Phaseolus vulgaris (common bean)||Fabaceae||Unknown|
|Prunus persica (peach)||Rosaceae||Unknown|
|Sorghum bicolor (sorghum)||Poaceae||Main|
|stored products (dried stored products)||Main|
|Triticum aestivum (wheat)||Poaceae||Unknown|
|Vigna angularis (adzuki bean)||Fabaceae||Unknown|
|Vigna unguiculata (cowpea)||Fabaceae||Unknown|
|Zea mays (maize)||Poaceae||Main|
Growth StagesTop of page
SymptomsTop of page
List of Symptoms/SignsTop of page
|Seeds / internal feeding|
Biology and EcologyTop of page
The biology of S. zeamais and S. oryzae has been reviewed in detail by Longstaff (1981). The adults are long-lived (several months to one year). Eggs are laid throughout most of the adult life, although 50% may be laid in the first 4-5 weeks; each female may lay up to 150 eggs. The eggs are laid individually in small cavities chewed into cereal grains by the female; each cavity is sealed, thus protecting the egg, by a waxy secretion (usually referred to as an 'egg-plug') produced by the female. The incubation period of the egg is about 6 days at 25°C (Howe, 1952). Eggs are laid at temperatures between 15 and 35°C (with an optimum around 25°C) and at grain moisture contents over 10%; however, rates of oviposition are very low below 20°C or above 32°C, and below about 12% moisture content (Birch, 1944).
Upon hatching, the larva begins to feed inside the grain, excavating a tunnel as it develops. There are four larval instars: in English wheat at 25°C and 70% RH, pupation occurs after about 25 days, although development periods are extremely protracted at low temperatures (e.g. 98 days at 18°C and 70% RH). Pupation takes place within the grain; the newly developed adult chews its way out, leaving a large, characteristic emergence hole. Total development periods range from about 35 days under optimal conditions to over 110 days in unfavourable conditions (Birch, 1944; Howe, 1952). The actual length of the life cycle also depends upon the type and quality of grain being infested: for example, in different varieties of maize, mean development periods of S. zeamais at 27°C and 70% RH have been shown to vary from 31 to 37 days. The development of S. zeamais on different wheats (Triticum spelta, T. dicoccum and T. monococcum spikelets and kernels) has also been studied (Suss et al., 1999). A demographic population simulation model of S. zeamais in grain stores in West Africa has been devised (Meikle et al., 1999).
Although both species are capable of flight, S. zeamais has a greater ability and tendency to fly (Giles, 1969). Where grain is stored on small farms, S. zeamais is thus more likely than S. oryzae to fly to the ripening crop in the field and establish an infestation in the grain before harvest.
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
|Acaropsellina docta||Predator||Arthropods|Larvae; Arthropods|Pupae|
|Anisopteromalus calandrae||Parasite||Arthropods|Larvae; Arthropods|Pupae|
|Cerocephala oryzae||Parasite||Arthropods|Larvae; Arthropods|Pupae|
|Lariophagus distinguendus||Parasite||Arthropods|Larvae; Arthropods|Pupae|
|Theocolax elegans||Parasite||Arthropods|Larvae; Arthropods|Pupae|
Notes on Natural EnemiesTop of page
ImpactTop of page
In stored maize, heavy infestations of these pests may cause weight losses of as much as 30-40%, although losses are commonly 4-5%. S. zeamais has been found to be amongst the most important pests of maize in a number of studies; in South Carolina, USA, (Arbogast and Throne, 1997), in Kentucky (Sedlacek et al., 1998) and in steel silos in Taiwan (Peng, 1998), and in central Italy (Trematerra et al., 1999). Insect pests infesting stored pearl millet and their damage potential, were assessed in northeastern Nigeria by Lale and Yusuf (2000). S. zeamais caused ca. 0.6% damage. Damage was greater in grains stored in the underground pit storage system than in grains stored in a rumbu, clay pot or polypropylene sack.
S. zeamais has been detected in seeds during quarantine processing of germplasm imported into India (Babu, 1997), in pigeonpeas imported from Indonesia and sorghum from the USA.
Infestations in rice are damaging, but loss estimates from real, as opposed to laboratory situations, are lacking.
Detection and InspectionTop of page
A male-produced aggregation pheromone which attracts both sexes occurs in Sitophilus spp. Sitophilure, 5-hydroxy-4-methyl-3-heptanone, was reported to be the aggregation pheromone common for S. oryzae and S. zeamais, and Levinson et al. (1990) confirmed the activity of 4S,5R sitophinone and 2S,3R-sitophilate for Sitophilus spp. Some interspecific cross attraction between three Sitophilus species has been demonstrated to occur; more so between S. oryzae and S. zeamais than between S. granarius and either of the other species.
The responses of S. zeamais to pheromone and synthetic maize volatiles as lures in crevice or flight traps have been studied in Kenya (Hodges et al., 1998). Three trap types (probe, cone and sticky) were used to monitor S. zeamais populations infesting shelled maize housed in galvanized steel storage bins. The results of this study were similar to those reported by Weston and Barney (1998); a combination of sticky traps in the grain bin headspace and probe traps positioned just below the grain surface was most efficient for monitoring the presence of pest and beneficial insect species in grain storage. Likhayo and Hodges (2000) reported the field monitoring of S. zeamais and S. oryzae using refuge and flight traps baited with synthetic pheromone and cracked wheat. The combination of pheromone and cracked wheat had an additive effect on trap catch.
Similarities to Other Species/ConditionsTop of page
Prevention and ControlTop of page
Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.
Cultural Control and Sanitary Methods
Good store hygiene plays an important role in limiting infestation by S. oryzae and S. zeamais. The removal of infested residues from last season's harvest is essential. The use of resistant maize varieties has also shown some potential in slowing the build-up of insect densities in stores. Phenolic compounds have been associated with grain resistance to S. zeamais.
A combination of controlled ambient aeration in the autumn and chilled aeration during summer storage has significant potential as a non-chemical preventative pest management technique (Maier et al., 1996). Aeration dramatically reduces the numbers of weevils compared with population levels in non-aerated maize (Arthur, 1998). Thorpe (1997) modelled ecosystems in ventilated conical-bottomed farm grain silos.
Zea mays genotypes resistant to S. zeamais have been evaluated by Boica et al. (1997) and in the field in Benin (Meikle et al., 1998). Genetic variability for resistance to S. zeamais in domestic US maize germplasm has been identified by Li et al. (1998). In Brazil, tests were carried out with maize cultivars to evaluate the attractiveness and oviposition preference of S. zeamais (Toscano et al., 1999).
The mechanism of resistance in maize to S. zeamais was investigated in relation to secondary chemistry and other biochemical and physical characteristics of maize genotypes. Phenolic acid content was correlated strongly with hardness of the grain (Arnason et al., 1997). Huang et al. (1998) assessed alpha-Pinene for antifeedant and growth inhibitory effects against S. zeamais adults.
Grain may be protected by the admixture of insecticide. Sitophilus spp. have a low susceptibility to synthetic pyrethroids but are readily killed by organophosphorous compounds such as fenitrothion and pirimiphos-methyl. Grain stocks may be fumigated with phosphine to eliminate existing infestation, but these treatments provide no protection against re-infestation. Sitophilus spp., particularly in the pupal stage, have a lower natural susceptibility to the fumigant phosphine and to carbon dioxide used in controlled-atmosphere storage than do other species tested and thus inadequate treatments are particularly likely to result in some survival.
Guedes and Heyde (1996) estimated the toxicity of deltamethrin to a number of resistant and susceptible populations of S. zeamais in traditional maize stores in Togo. In another study, a treatment with pirimiphos-methyl and deltamethrin was most effective in traditional granaries after pirimiphos-methyl, deltamethrin and permethrin were applied in various combinations (Richter et al., 1997, 1998). Srinivasacharayulu and Yadav (1997) evaluated the toxicity of deltamethrin, fluvalinate, chlorpyrifos-methyl, etrimfos and malathion against S. zeamais, and topical application bioassays of malathion, pirimiphos-methyl, deltamethrin and permethrin were undertaken with 11 field strains of S. zeamais, collected from nine states in Mexico (Perez-Mendoza, 1999).
A number of studies have investigated the efficacy of phosphine fumigation: Goto et al. (1996) reported its effects at concentrations of 0.5, 1 or 2 g/m³ on pupae at 15°C. The effects of phosphine on the development of S. zeamais was investigated by Zhang (1997). Field strains of S. zeamais from the Philippines were evaluated for resistance to phosphine by Gibe et al. (1997). Peng et al. (1999) and Peng (2000) studied vertical penetration and distribution of phosphine in corn and sorghum stored in steel silos and compared the distribution of phosphine in bins when aluminium phosphide was placed on the stack surface and on the floor.
Guedes et al. (1996) reported that 20% carbon dioxide controlled S. zeamais in 5 days. Treatment with 15% carbon dioxide and 5% oxygen controlled the insects in 10 days. Santos et al. (1997) describes use of carbon dioxide-controlled atmosphere in Brazil for the control of S. zeamais on maize and wheat. Carbon dioxide at concentrations of 50% and 60% killed all life stages of the weevil after 10 days of exposure. In another study with fumigation periods of 15 and 20 days, the concentrations of 40%, 50% and 60% eliminated all stages of the insects (Santos et al., 1998). According to Casella et al. (1998), effective control of Sitophilus was achieved with a synthetic atmosphere and 0.5 or 0.75 g/m³ of phosphine, for an exposure period of 120 hours.
Natural carbon dioxide production in stored maize is affected by moisture content, the amount of broken corn and foreign materials and infestation by S. zeamais (Sone, 1999). Respiration by fungi including Aspergillus spp. appears to have a greater influence on carbon dioxide production in stores than the presence of insects. Under sealed storage conditions in maize, insects and fungi both deplete the oxygen supply, creating an unfavourable atmosphere for their own survival (Damcevski et al., 1998; Moreno-Martinez et al., 2000).
Contessi (1999) assessed the effectiveness of controlled atmospheres using generators of inert gases, such as carbonic anhydride and nitrogen, for the disinfestation of wheat stored in vertical silos and horizontal stores.
Irradiation of S. zeamais by microwave and gamma radiation has been studied. Insect mortality studies were carried out with a high-power microwave source operating at a frequency of 10.6 GHz at power levels of 9-20 kW to irradiate samples of Triticum aestivum infested with S. zeamais (Halverson et al., 1996). Applying high-power microwaves demonstrated that selective heating of insects and their resulting mortality is a non-linear function of frequency and increases at frequencies above 2.45 GHz in the vicinity of increasing relaxation processes associated with free water (Plarre et al., 1997). Tests were conducted at 12, 15, 17.9 and 55 GHz. The results indicated that 15 GHz produced greater mortality among adults than 12 and 17.9 GHz. However, 55 GHz produced the greatest mortality for the three developmental stages studied.
A sterilizing dose of gamma radiation from Cobalt-60 was determined for adults of S. zeamais on rice, maize and wheat grains (Franco et al., 1997) at 24-26°C and 65-75% RH with a dose rate of 3.00 kGy/h. The dose was found to be 60 Gy for S. zeamais.
Low-temperature storage of rice is extensively practised to control insect pests in Japan. Nakakita et al. (1997) reported that both hatching and metamorphosis of S. zeamais were completely inhibited at 10°C; a small number of adult S. zeamais emerged at 15°C.
Dry heat treatment has been found to be an effective control against all developmental stages of S. zeamais (Mohammed-Dawd and Morallo-Rejesus, 2000). All eggs and adult weevils were killed following exposure to 60°C for 2 hours, or 70-80°C for 1 hour. All larvae were killed when seeds were exposed to 70-80°C for 1 hour.
The bionomics of the pteromalid parasitoid Lariophagus distinguendus and its controlling effect on S. zeamais were studied by Li et al. (1998). L. distinguendus was found to have five generations each year in the laboratory, with final-instar larvae of the parasitoid over-wintering in larvae of S. zeamais. The parasitoid Theocolax elegans has also been recorded to attack S. zeamais (Helbig, 1998).
The effect of different isolates and formulations of Beauveria bassiana on S. zeamais in stored maize are reported by Adane et al. (1996); Moino and Alves (1997); Hidalgo et al. (1998); and Junior and Alves (1998). B. bassiana can be an effective microbial control agent if used as a preventative treatment (Moino et al., 1998).
The fungal pathogens of maize storage pests in Kenya were surveyed by Oduor et al. (2000). Between 0.08 and 0.94% of the insects collected (mostly S. zeamais) were found to be infected.
The toxicity of a large number of essential oils and plant parts of various spices and herbs have been assessed against S. zeamais.
A number of citrus oils have vapour toxicity to adults of S. zeamais; lime, mandarin and grapefruit peel oils reduce oviposition and larval emergence (Don-Pedro, 1996). Garlic oil is toxic to adults (Ho et al., 1997), and oils of coconut, sunflower, sesame and mustard, alone and in combination with eucalyptol, eugenol or camphor have been found to be toxic to S. zeamais in wheat and maize-treated grains (Obeng-Ofori and Reichmuth, 1999).
Plants or extacts from a considerable number of other plants have exhibited toxicity to S. zeamais. These include: Ocimum suave (Bekele et al., 1996); Ocimum kenyense (Bekele et al., 1997); Ocimum kilimandscharicum (Obeng-Ofori et al., 1998); Dennettia tripetala, Piper guineense, Monodora myristica and Xylopia aethiopica (Okonkwo and Okoye, 1996); Ricinus communis, Gaura coccinea, Larrea tridentata, Ribes ciliatum, Castilleja tenuiflora, Alchemilla procumbens and Guazuma tomentosa (Araya-Gonzalez et al., 1996); Juniperus sabina (Gao and Zhang, 1997); Chrysanthemum cinerariaefolium (Saggar et al., 1997); Tagetes minuta (Weaver et al., 1997); cinnamon, garlic, coriander, Capsicum annuum and Murraya paniculata (Ho et al., 1997); Eucalyptus (Obeng-Ofori et al., 1997); Cinnamomum aromaticum (Huang and Ho, 1998); Pachyrhizus erosus (Alavez-Solano et al., 1998); Allium sativum (Chiam et al., 1999); Evodia rutaecarpa (Liu and Ho, 1999); Canaga odorata (Huang et al., 1999); Baccharis genistelloides (Prates, 1999); Aloe marlothii (Achiano et al., 1999); Securidaca longipedunculata (Belmain et al., 1999); Cleome hirta; cardamom (Huang et al., 2000); Basella alba, Operculina turpethum and Calotropis gigantea (Haque et al., 2000).
ReferencesTop of page
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