Rhyzopertha dominica (lesser grain borer)
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- Hosts/Species 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
- Links to Websites
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Rhyzopertha dominica (Fabricius)
Preferred Common Name
- lesser grain borer
Other Scientific Names
- Apate pusilla Fairmaire 1850
- Apate rufa Hope 1845-47
- Bostrychus moderatus Walk.
- Dinoderus frumentarius Motschulsky 1857
- Dinoderus pusillus Horn 1878
- Ptinus fissicornis Marsham 1802
- Ptinus picus Marsham 1802
- Rhizoperta dominica (F.)
- Rhizopertha dominica Lesne 1896
- Rhizopertha pusilla Stephens 1830
- Rhizopertha rufa Waterhouse 1888
- Rhyzopertha pusilla Fabricius
- Synodendron dominica Fabricius
- Synodendron dominicum Fabricius 1792
- Synodendron pusillum Fabricius 1798
International Common Names
- English: American wheat weevil; grain, borer, lesser; grain, borer, stored; grain, eater, small; weevil, Australian wheat
- Spanish: barrenador grande de los granos; barrenador menor de los granos; capuchino de los granos; escarabajo de los granos; gorgojo da los creales; gorgojo de los cereales; pequeño barrenador del trigo; taladrillo de los granos
- French: bostryche des grains; capucin des grains; perceur (petit) des céréales; perceur des cereales, petit; petit perceur des céréales
- Portuguese: besourinho do trigo armazenado
Local Common Names
- Czechoslovakia (former): korovník obilní
- Germany: Getreidekapuziner
- Hungary: Gabonaalszu
- Indonesia: Gabah-bubuk
- Israel: norer hatiras
- Italy: punteruolo dei cereali
- Netherlands: Graanboorder, kleine
- Norway: kornborer
- Poland: kapturnik zbozowiec
- Turkey: ekin kambur biti
- Yugoslavia (Serbia and Montenegro): kapuciner; rizoperta; zitni kukuljicar
- RHITDO (Rhyzopertha dominica)
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Arthropoda
- Subphylum: Uniramia
- Class: Insecta
- Order: Coleoptera
- Family: Bostrichidae
- Genus: Rhyzopertha
- Species: Rhyzopertha dominica
Notes on Taxonomy and NomenclatureTop of page See Gardner (1933) for a key separating full-grown larvae of the family Bostrichidae.
DescriptionTop of page Potter (1935) provided a detailed description of all life stages of R. dominica. The egg is typically white when first laid, turning rose to brown before hatching. The egg is ovoid in shape, 0.6 mm in length, 0.2 mm in diameter. There are usually four larval instars. The larvae are scarabaeiform, the first two instars are not recurved, the third and fourth instars have the head and thorax recurved towards the abdomen. The widths of the head from the first to the fourth instar are 0.13, 0.17, 0.26 and 0.41 mm, and the lengths of the larvae are 0.78, 1.08, 2.04 and 3.07 mm, respectively. The pupae are 3.91 mm in length, with 0.7 mm between the eyes. At the end of the abdomen, the male pupae have a pair of 2-segmented papillae fused to the abdomen for their entire length, whereas female papillae are 3-segmented and project from the abdomen. Adults are 2-3 mm in length, reddish-brown and cylindrical. The elytra are parallel-sided, the head is not visible from above, and the pronotum has rasp-like teeth at the front.
DistributionTop of page R. dominica is thought to originate from the Indian subcontinent, but now has a cosmopolitan distribution. It is a serious pest of stored products throughout the tropics, Australia and the USA. It is also found in temperate countries, either because of its ability for prolonged flight or as a result of the international trade in food products.
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.
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Bangladesh||Widespread||Taylor and Halliday, 1986|
|Bhutan||Present||Taylor and Halliday, 1986|
|China||Present||Dunkel et al., 1982|
|India||Widespread||Sinha and Sinha, 1990|
|-Andhra Pradesh||Present||Reddy and Swamy, 2006|
|-Assam||Present||Sikha Deka, 2003|
|-Bihar||Present||Sinha and Sinha, 1990|
|-Delhi||Present||Bera et al., 2007|
|-Himachal Pradesh||Present||Ramesh et al., 2000|
|-Indian Punjab||Present||Dhaliwal, 1977|
|-Karnataka||Present||Hampanna et al., 2006|
|-Odisha||Present||Sahu et al., 2008|
|-Rajasthan||Present||Balbir et al., 2003|
|-Tamil Nadu||Present||Kumar et al., 2005|
|-Uttar Pradesh||Present||Girish et al., 1974|
|Indonesia||Present||Soegiarto et al., 1981; Sidik et al., 1985|
|-Java||Present||Soegiarto et al., 1981|
|Iran||Present||Khormaly et al., 2002|
|Iraq||Present||Ismail et al., 1988|
|Israel||Present||Carmi and Pater, 1989|
|Korea, DPR||Present||Paik, 1976|
|Malaysia||Present||Seth, 1975; Muda, 1985|
|Nepal||Present||Taylor and Halliday, 1986|
|Pakistan||Present||Tilton et al., 1983; Taylor and Halliday, 1986|
|Philippines||Present||Caliboso et al., 1985; Sayaboc and Acda, 1990|
|Saudi Arabia||Present||Mostafa et al., 1981; Rostom, 1993|
|Sri Lanka||Present||Ganesalingam, 1977; Taylor and Halliday, 1986|
|Syria||Present||Teriaki and Verner, 1975|
|Taiwan||Present||Lin, 1981; Lo, 1986; Lin et al., 1990|
|Turkey||Present||Aydin and Soran, 1987; Yucel, 1988|
|Vietnam||Present||Stusak et al., 1986|
|Algeria||Present||Mebarkia et al., 2009|
|Chad||Present||Trematerra et al., 2003|
|Egypt||Present||El Nahal et al., 1984|
|Ghana||Present||Belmain et al., 2001|
|Mali||Present||Taylor and Halliday, 1986|
|Morocco||Present||Bartali et al., 1990|
|Namibia||Present||Stejskal et al., 2006|
|Nigeria||Present||Ivbijaro, 1979; Ekundayo, 1988|
|Rwanda||Present||Weaver et al., 1991|
|South Africa||Present||Viljoen et al., 1984|
|Tanzania||Present||Hodges et al., 1983|
|Zimbabwe||Present||Mvumi et al., 2003|
|Canada||Present||Fields et al., 1993|
|-Alberta||Present||Fields et al., 1993|
|-British Columbia||Present||Fields et al., 1993|
|-Manitoba||Restricted distribution||Fields et al., 1993|
|-New Brunswick||Present||Fields et al., 1993|
|-Quebec||Restricted distribution||Fields et al., 1993|
|-Saskatchewan||Present||Fields et al., 1993|
|Mexico||Present||Rojas, 1988; Corral et al., 1992|
|USA||Widespread||Storey et al., 1982; Storey et al., 1983|
|-Montana||Present||Watts and Dunkel, 2003|
|-Oklahoma||Present||Mahroof and Phillips, 2006|
|-Oregon||Present||Cuperus et al., 1990|
|-Texas||Present||Arthur et al., 2008|
Central America and Caribbean
|Cuba||Present||Aviles and Guibert, 1986|
|Puerto Rico||Present||Potter, 1935|
|Brazil||Present||Taylor and Halliday, 1986; Pacheco et al., 1990|
|-Espirito Santo||Present||Botelho and Arthur, 2001|
|-Rio Grande do Sul||Present||Oliveira et al., 1990|
|-Santa Catarina||Present||Trematerra et al., 2004|
|-Sao Paulo||Present||Pacheco et al., 1990|
|Former USSR||Present||Asanov, 1980|
|Germany||Present||Bahr, 1975; Rassmann, 1978|
|Italy||Present||Trematerra and Daolio, 1990; Suss et al., 1991|
|Poland||Present||Klejdysz and Nawrot, 2010|
|Russian Federation||Present||Zakladony, 1990; Asanov, 1980|
|-Russia (Europe)||Present||Podobivskii, 1991|
|Spain||Present||Pascual-Villalobos et al., 2006|
|Switzerland||Present||Hoppe, 1981; Buchi, 1993|
|UK||Present||Dyte et al., 1975; Jacobson and Thomas, 1981|
|Yugoslavia (former)||Present||Pireva, 1992|
|Australia||Widespread||Sinclair and Haddrell, 1985; Collins et al., 1993|
|-New South Wales||Widespread||Greening, 1979|
|-Victoria||Present||Daglish et al., 2008|
HabitatTop of page Rhyzopertha dominica is found mainly in cereal stores, and food and animal feed processing facilities. It has also been trapped using pheromone-baited flight traps several kilometres from any food storage or processing facility (Fields et al., 1993).
Hosts/Species AffectedTop of page Adults and larvae of R. dominica feed primarily on stored cereal seed including wheat, maize, rice, oats, barley, sorghum and millet. They are also found on a wide variety of foodstuffs including beans, dried chillies, turmeric, coriander, ginger, cassava chips, biscuits and wheat flour. There are several reports of the lesser grain borer being found in or attacking wood (Potter, 1935), as is typical of other Bostrichidae. R. dominica has been reported to produce progeny on the seeds of some trees and shrubs (acorns, hackberry [Celtis occidentalis] and buckbrush [Symphoricarpos orbiculatus]) (Wright et al., 1990).
Growth StagesTop of page Post-harvest
List of Symptoms/SignsTop of page
|Seeds / external feeding|
Biology and EcologyTop of page Females of R. dominica lay between 200 and 500 eggs in their lifetime. The eggs are laid loose in grain. The lowest temperature at which R. dominica can complete development is 20°C; at this temperature, the development from egg to adult takes 90 days. The fastest rate of development occurs at 34°C; at this temperature the egg takes 2 days, the larvae 17 days, and the pupae 3 days to complete development. R. dominica is unable to complete development between 38 and 40°C. Adults live for 4-8 months. Under optimal conditions of 34°C and 14% grain moisture content, there is a 20-fold increase in the population of R. dominica after 4 weeks. It can successfully infest grain at 9% moisture content, but has higher fecundity, a faster rate of development, and lower mortality on grain of a higher moisture content.
Adult males produce an aggregation pheromone in the frass that attracts both male and female adults. Adults are good flyers, and can be trapped in pheromone-baited flight traps placed several kilometres from grain stores. Adults can bore into intact kernels. The larvae of R. dominica are mobile.
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
Notes on Natural EnemiesTop of page There are a few predators of R. dominica. Teretriosoma nigrescens is a histerid beetle that is found in Central America feeding on Prostephanus truncatus. It also feeds on R. dominica, though it produces more offspring on P. truncatus (Puschko, 1994). This predator was released into Africa in 1991 in an effort to control P. truncatus, and is now well established in Togo and Benin in West Africa, and in Kenya in East Africa. However, the ability of T. nigrescens to significantly reduce P. truncatus or R. dominica populations has yet to be determined (Markham et al., 1994). There is some concern regarding the use of T. nigrescens, as it also feeds on grain. The cadelle Tenebroides mauritanicus also feeds on grain, mites and stored-product insect eggs, including Rhyzopertha (Bousquet, 1990; Yoshida, 1975). The predatory mites, Cheyletus eruditus and Pyemotes ventricosus feed on a wide variety of stored product insect eggs (Asanov, 1980; Brower et al., 1991), but their effect on populations in the field has not been determined. The anthocorid bugs, Xylocoris flavipes and Lyctocoris campestris, are general predators that are known to attack R. dominica and are found in grain stores around the world (Parajukee and Phillips, 1993; Asanov, 1980).
ImpactTop of page
R. dominica is a major pest of wheat (Flinn et al., 2004) and rice (Chanbang et al., 2008a,b) around the world. Both larvae and adult produce frass and cause weight losses by feeding on grains. R. dominica infestation can reduce rice to dust (Emery and Nayak, 2007).
There are three aspects of the impact of R. dominica infestation: loss in the quantity of stored grain, loss in quality of stored seeds (Sánchez-Mariñez et al., 1997) and the cost to prevent or control infestations (Cuperus et al., 1990; Anonymous, 1998).
On wheat and rice, larvae consume both germ and endosperm during their development in grain and thus produce more frass than Cryptolestes ferrugineus and Sitophilus granarius (Campbell and Sinha, 1976). R. dominica is also capable of damaging grain, causing weight losses of up to 40%, compared to 19%, 14% and 10% for S. oryzae, Tribolium castaneum and Ephestia cautella, respectively (Sittusuang and Imura, 1987). Weight loss from individual kernels has also been reported with different varieties of triticale, a wheat-rye hybrid (Baker et al., 1991), and in rice infested with R. dominica (Nigam et al., 1977). R. dominica feeding on seed germ reduces germination rates and vigour of the grains and may be followed by secondary pests and fungi (Bashir, 2002).
Food production and nutritional value
R. dominica infestation of wheat, maize and sorghum grains resulted in substantial changes in the contents of calcium, phosphorus, zinc, iron, copper and manganese (Jood et al., 1992). Jood and Kapoor (1992) also observed a reduction in the starch digestibility of maize, rice and sorghum in response to R. dominica infestation. Single or mixed populations of Trogoderma granarium (Khapra beetle) and R. dominica resulted in substantial reductions in the contents of total lipids, phospholipids, galactolipids and polar and nonpolar lipids of wheat, maize and sorghum (Jood et al., 1996). R. dominica has also been reported to decrease vitamin contents of grain; 75% level of infestation of cereal grains caused losses of 23 to 29% (thiamine), 13 to 18% (riboflavin) and 4 to 14% (niacin) we (Jood and Kapoor, 1994).
Chapatis prepared from flours with more that 50% R. dominica and T. granarium infestation level tasted bitter (Jood et al., 1993). At 75% infestation level there was a significant reduction in protein nitrogen and true protein contents of three cereal grains (Jood and Kapoor, 1992).
It is difficult to estimate the actual costs incurred for the control of R. dominica because it is generally found in mixed population with other stored-product insect pests that also cause damage. The species R. dominica associates with vary depending upon the region and stored commodity. Two or more live ‘grain-damaging’ insects per kg of wheat resulted in an infested designation on the grain inspection certificate (FGIS, 1997). R. dominica produces insect-damaged kernels (IDK) when adults emerge from the kernels. If wheat contains more than 32 IDK per 100 g it is designated as sample grade, which cannot be sold for human consumption, and its market value drops dramatically (FGIS, 1997).
Laboratory experiments have estimated that one R. dominica consumes 0.15 g of wheat in its life time (Campbell and Sinha, 1976; Storey et al., 1982). If all R. dominica completed their life cycle from 1976 to 1979 in the USA, they would have consumed 300,000 metric tonnes of wheat annually, or 0.5% of the total of stored wheat. Similar calculations, assuming R. dominica eats 0.15 g during its lifetime, give 8000 tonnes of maize (0.004% loss of total harvest) and 2000 tonnes of oats (0.02% loss) consumed annually by R. dominica.
Data collected from 1998 to 2002 from wheat stored in commercial grain elevators in south-central Kansas, USA, revealed that C. ferrugineus, R. dominica and T. castaneum were the primary insect species found in collected wheat samples. In the top 3.7 m of grain, R. dominica made up 44% of the insects found in the samples, and from 3.8 to 12.2 m it was present at 84% (Flinn et al., 2010). In 2008, R. dominica caused serious damage to grain in an Indonesia government storage unit (Astuti et al., 2013).
The cost of controlling storage pests can be substantial. The Environmental Protection Agency in the USA estimated that 270,000 kg of aluminum phosphide pesticide was used annually between 1987 and 1996 for wheat in storage, roughly one third of the total aluminium phosphide used in those years (Anonymous, 1998).
DiagnosisTop of page See section on Detection and Inspection Methods.
Detection and InspectionTop of page A variety of methods have been used to detect insect pests of stored products, including R. dominica. The simplest method is to sieve a 200-1000 g sample of the grain and look for adults. However, only a small sample of the grain is inspected using this method and larvae, pupae and adults inside the grain kernels are not detected. An inclined sieve enables 30 kg of grain to be sampled in 5 minutes, with the extraction of 90% of the adults (White, 1983). Berlese funnel extractions can be used to extract adults and larvae from inside the seed, by forcing the insects out of the grain into collection jars. This method does not detect non-mobile stages such as the pupae and eggs; it also takes several hours and requires specialized equipment.
An ELISA test which detects the presence of the insect muscle protein, myosin, can also be used (Quinn et al., 1992). The ELISA test is mainly intended for use in flour mills to ensure that the flour does not contain large numbers of insect fragments or whole insects, but it can also be used to detect insects in whole grain. The cost of the ELISA and the time involved make it unlikely to replace Berlese funnel extractions. However, rapid and inexpensive ELISA kits are being developed for use in the field and may become available in the near future.
Lesser grain borer can be detected by placing probe pitfall traps in the grain (White et al., 1990; Hagstrum, 2000). Insects fall into these traps, which resemble a torpedo with holes, as they move through the grain. The traps are left in the grain and inspected periodically. This method is more sensitive than the extraction of insects using either a sieve or Berlese funnel. However, the traps should be left in place for 2-7 days and are ineffective at temperatures below 10°C, when the insects stop moving. Placing the trap in the grain can also be impractical in many storage situations. To address this limitation, grain probe traps have been wired to electronic sensors that detect insects as they fall into the trap (Epsky and Shuman, 2001). Pheromone-baited flight traps are a highly sensitive tool for the detection of R. dominica (Leos Martinez et al., 1987; Fields et al., 1993). Like probe pitfall traps, flight trap catches are a relative measure and are temperature dependant, with few insects being caught at air temperatures below 20°C.
Near-infrared spectroscopy (NIRS) has been used for many years commercially to measure protein content in grain. In the laboratory, NIRS can be used to detect late instar larvae, pupae and adults of R. dominica and other internal feeders in grain (Dowell et al., 1998). This system can also be used to distinguish R. dominica from other insects (Dowell et al., 1999).
Temperature, carbon dioxide, sound and feeding damage can be used as indirect indicators of insect infestation. Insects release heat as they respire, and at high insect densities, 'hot spots' can be created. Thermometers, either attached to the end of a probe or wired permanently into a storage structure with a remote electronic readout, give a quick and accurate picture of grain temperature. However, hot spots can be very localized and difficult to detect, and insects can cause significant damage before they are detected.
Increased carbon dioxide concentration in the grain indicates the presence of insects in the storage bin at an earlier stage, during the development of the insect infestation. This method allows detection at lower insect densities, at the beginning of an infestation. Tubes are placed into the top of the grain mass, in the centre, and the intergranular air is sampled every 2 weeks (Sinha et al., 1986). Both of these methods also detect the growth of moulds.
Researchers have been listening to insects in stored grain for over 40 years (Shade et al., 1990; Hagstrum et al., 1990). Commercial units are now available to detect the sounds of insects moving and feeding. This technology is in its infancy and a number of improvements are needed before it can become an effective tool for the detection of insect infestations.
Feeding holes in grain and other commodities also indicate the presence of some insects, but sampling using sieves or traps is needed to identify which insects have caused the damage. All of these detection methods are not specific for R. dominica, and grain samples are required for verification of the insect species.
Similarities to Other Species/ConditionsTop of page The Bostrichidae can be distinguished from most other Coleoptera found in stored foodstuffs by the presence of rasp-like teeth on the pronotum. Some Scolytidae found in stored products also have pronoted aspirities but but with a compact club not loose 3-segmented as in the Bostrichidae.
R. dominica can be distinguished from Prostephanus truncatus, the larger grain borer, by the shape of the abdomen and the elytra. The posterior end of the abdomen of Prostephanus is square with distinct corners, whereas Rhyzopertha has a tapered abdomen. When viewed from the side, the elytra declivity, where the elytra slopes towards the tip, is steep and flat for Prostephanus and tapered and rounded for Rhyzopertha. Less reliable indicators are adult size and colour. Prostephanus is black and 3-4 mm in length, whereas Rhyzopertha is brown and 2-3 mm in length.
In addition, the following characters may also be used to distinguish between these species.
R. dominica has pronotum with coarse asperities anteriorly, less coarse granules posteriorly, anterior margin strongly dentate in a regular curve. Elytra with regular rows of coarse punctures (finer at sides) covered with curved setae, fine anteriorly, thick posteriorly: declivity unarmed.
P. truncatus has pronotum with coarse asperities on front half (anterior - most ones forming inverted V which projects at middle of front margin), with fine granules on posterioir half which change to single punctures at sides. Elytra with coarse, partly linear punctation, covered with long, fine setae, curved anteriorly, erect posteriorly; declivity with lateral and preapical carina, and slightly raised suture.
Another bostrichid found in stored products, Dinoderus minutus, the bamboo powderpost beetle, can be distinguished from Rhyzopertha by its stout body and a pair of shallow, medial depressions near the base of the pronotum (Bousquet, 1990). Also, the elytra are irregularly punctate and declivity with ocellate punctures in Dinodermus. D. minutus is a pest of bamboo and cane which is occasionally found in stored grain, tobacco and fruit.
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.
Physical control of R. dominica involves the manipulation of the temperature, relative humidity, atmospheric composition (air gases composition), sanitation, ionizing radiation and the removal of adult insects from the grain either by sieving or air. All these practices may be helpful in eliminating or reducing insect pest infestations to a tolerable level (Jayas et al., 1995).
Sanitation and hermetic sealing
Cleaning during grading operations, drying, cool storage and hermetically sealed packaging can all play an effective role in conserving the seed viability with residue free pest control.
Grain packaging in airtight structures is one of the most important physical methods controlling R. dominica. These structures may range from well-sealed barrels holding several kilograms to 100-t capacity metal bins. The structures should be pressure-tested to confirm airtightness. Portable hermetic storage bags are also available (Garcia-Lara et al., 2013).
Removing insects by sieving is not equally effective for all species as several insect species, including R. dominica, spend most time of their life cycle remaining inside the grain or kernel. Impacting the grain, either by moving the grain using a pneumatic conveyer or dropping the grain onto a spinning, studded disc, can reduce R. dominica populations by over 90%. Good sanitation, particularly the removal of spilt grain around storage facilities, is a preliminary step in reducing insect populations that can infest grain in storage.
Aeration and drying
One of the more effective non-chemical control methods is to cool the grain with aeration fans, which gradually suppresses insect population growth in the storage period. The Kansas State extension program has advocated early aeration as the best non-chemical insect suppression method; a study conducted in Kansas, USA, exhibited that aeration starting from harvesting, using automatic fan controllers, allowed safe storage of grain for several months (Reed and Harner, 1997).
A moisture content (mc) of 25% is not uncommon in newly harvested grain in humid regions, but grains with 14% mc can be safely stored for 2-3 months. For longer storage periods, from 4-12 months, the moisture content must be reduced further. Reducing grain moisture content reduces the number of eggs produced and the survival of offspring and adults. There are 3 types of drying: ambient air drying, sun drying and mechanical drying. In ambient air drying system, air is heated and passed through grain to produce a relatively high vapour pressure gradient between the moisture in the grain and the moisture in the drying air. This gradient causes moisture to move from the grain into the air, where it is then exhausted from the grain bulk to the outside atmosphere. (Jones et al., 2012). In many countries in Asia, Africa, and Latin America grain drying is achieved by spreading a thin layer of grain in the sun, on the threshing floor or on rooftops. A mechanical way to remove the water from wet grains is by blowing (heated) air through the grain. Mechanical drying of wheat grain is not practiced in many the developing countries, which largely rely on sun drying. At 34°C and 14% moisture content there were 109 R. dominica adults produced per female per generation; 10 adults at 10% moisture content, 0.3 adults at 9% and none at 8% moisture content (Birch, 1953).
Radio-frequency heat treatment is increasingly used as a new thermal method for the disinfection of post-harvest insect populations in agricultural commodities (Tang et al., 2000). The application of this method leaves no chemical residue and provides acceptable product quality with minimal environmental impacts (Wang et al., 2003). Janhang et al. (2005) evaluated the efficiency of radio-frequency heat treatment against R. dominica both on the seed surface and inside the seed. The rice kernels with 10.4% moisture content and 93% germination rate were treated with radio frequency heat treatment at 27.12 MHz at 70, 75, 80 and 85 °C for 180 seconds. 100% mortality of R. dominica was achieved in all treatments; however, the rice seed quality was also decreased at higher temperatures. Phosphine-resistant adults were found to be more tolerant than phosphine-susceptible adults toward soft-electron and gamma radiation (Hasan et al., 2006).
More recently, a flameless catalytic infrared emitter was used to disinfest hard red winter wheat containing different life stages (eggs, larvae, pupae and 2-week-old adults) of R. dominica (Khamis et al., 2010). Approximately 94% mortality of all R. dominica life stages occurred when 113.5 g of wheat was exposed for 60 s at a distance of 8.0 cm from the emitter, resulting in wheat temperatures that ranged between 107.6 ± 1.4 and 113.5 ± 0.5°C. These findings suggested this technology as a promising tool for disinfestation of stored wheat (Khamis et al., 2010).
Reducing temperatures to below 34°C reduces the rate at which the population of R. dominica increases. R. dominica cannot complete its life cycle below 20°C. In temperate countries, grain temperatures can be reduced by forcing air from outside through the grain, especially in winter. Grain can also be cooled by aeration using refrigerated air. Commercial units are available for both types of cooling. Increasing grain temperature to above 34°C also reduces the rate at which the population of R. dominica increases.
Although R. dominica is one of the most heat tolerant of all stored grain insect pests, it can be controlled by heating the grain to 65°C in 4 minutes, and rapidly cooling it to below 30°C. Commercial units that can handle 150 t of grain/h have been developed in Australia. The running costs of these units are comparable to those of chemical control. Care must be taken to ensure that the commodities in storage are only heated briefly so that the quality of the grain is not reduced. Heat disinfestation of grain has the potential for higher market acceptance than chemically treated grains.
Two interrelated ways have been used to make the method more affordable: one is to decrease typical grain treatment temperatures and hold these temperatures for a time sufficient for disinfestation; the other is to increase the rate of heating to induce physiological heat shocks, thereby bringing about faster insect mortality.
Using a spouted bed, the rates of heat-tolerant species R. dominica mortality were recorded over a range of grain temperatures (50°C to 60°C) at 12% moisture content. It was observed that when the initial rate of heating was increased by increasing the air inlet temperature from 80-100°C, the time required for a given level of mortality was significantly decreased. Moreover, decreasing the target grain temperature and increasing the treatment period accounted for additional cost savings. For instance, at the most rapid rate of heating, grain that reached 60°C required 0.73 min of heat soak for 99.9% mortality and cost a theoretical US$ 2.72/t, while grain that reached 55°C required 23.62 min for the same level of mortality and cost US$ 1.87/t. By 50°C, 22 h were required, but the theoretical running cost was reduced to US$ 1.25/t (Beckett and Morton, 2003). Manipulation of storage temperature is a relatively new technology that may be used to a greater extent in the future.
The manipulation of gases (nitrogen (N2), oxygen (O2) and carbon dioxide (CO2) within storage structures has been widely studied for the control of insect infestations. The two main approaches involve increasing CO2 concentration and reducing oxygen in the storage vicinity. To control the insect infestations, oxygen levels must be maintained below 1% for 20 days, or carbon dioxide levels maintained at 80% for 9 days, 60% for 11 days or 40% for 17 days. The storage structures should be sealed properly before the addition of gases (Annis and Graver, 1990).
The effectiveness of CO2 at different temperatures (20, 25, 30, 35 and 40°C) and exposure intervals (6, 12, 18, 24, 30, 36, 48, 54 hr) was tested against the life stages of different stored grain insects including R. dominica. The eggs of R. dominica were particularly tolerant at 20°C, which required extended exposure to treatment (54 and 48 hr, respectively) to prevent the egg survival. The adults were highly susceptible and a 24-hr exposure at 20°C or 6-hr at temperatures of >30°C were enough to achieve 100% mortality (Locatelli and Daolio, 1993). The combination application of carbon dioxide (5-20%) with the fumigant ethyl formate significantly enhanced the effectiveness of the fumigant against R. dominica and living stages of some other stored grain insect species (Haritos et al., 2006).
Inert dusts have been used as a traditional method of insect control for thousands of years (Glenn and Puterka, 2005). Stored grain insects are more vulnerable to these dusts as they feed upon dry grains and possess relatively larger surface to volume ratio (Stathers et al., 2004). There are several types of inert dusts being used in insect control programs, such as ash, lime, clay, diatomaceous earth (DE) and silica aerogel. The most effective inert dusts are DE and silica aerogel. Silica aerogels are man-made powders with smaller and more uniform particle sizes than DE. A major portion of silica aerogel and DE is made up of silicon dioxide, which dehydrates the insect body by both cuticle lipid absorption and abrasion (Quarles and Winn, 1996). Although DEs have low mammalian toxicity (Athanassiou et al., 2004), most DE formulations are used at considerably high application rates for the effective insect pest control (Vayias et al., 2006). At high concentrations, DE reduces grain bulk density by 9% and flow rate by 39% (Jackson and Webley, 1994), which is considered unacceptable for many large-scale commercial farms. These levels of loss may be more acceptable for households or subsistence farms.
DEs from different geological sources have different efficacies (Nwaubani et al., 2014), and the concentrations required to control infestations must be assessed before use. Other factors determining the efficacy of any DE formulation include: test insect population (Wakil et al., 2013), test insect species (Vassilakos et al., 2006), exposure interval (Baldassari et al., 2008), dose rate (Wakil et al., 2010) and temperature and relative humidity (Chanbang et al., 2007).
R. dominica is relatively tolerant to DE, and concentrations between 500 and 1000 ppm are required to control populations (Subramanyam et al., 1994). However, by the introduction of enhanced DE formulations, an effective control of this insect species is possible, even at a dose rate of 50 ppm (Wakil et al., 2011; Riasat et al., 2013). Furthermore, the combined use of DE with other insect control methods could provide effective control of R. dominica (Lord, 2005; Athanassiou et al., 2008; Riasat et al., 2011; Wakil et al., 2012; 2013).
Host plant resistance
Although there are substantial differences in the resistance of host varieties to R. dominica (Kishore, 1993; Cortez-Rocha et al., 1993), the use of resistant varieties has not been exploited as a method of control. Resistant varieties often do not prevent insect infestations, but reduce the rate at which infestations develop, and increase mortalities. Host resistance would enable the crop to be stored for a longer period before extensive damage is caused by insect populations. 28 different varieties of short, long and medium size rice kernel exhibited variable level of resistance to R. dominica, as measured on the Dobie Index of susceptibility (Chanbang et al., 2008b).
Adult R. dominica mortality due to DE treatment was generally greater on more resistant varieties of rough rice compared to less resistant varieties (Chanbang et al., 2008a).
The susceptibility of six milled rice varieties (IR-64, Ciherang, Membramo, Cibogo, Sembada, and Intani-2) to R. dominica was studied under laboratory conditions (Astuti et al., 2013). The susceptibility was measured on the basis of number of eggs laid by female insects, the number of F1 progeny emerged, the weight loss of infested samples and also using the Dobie index of susceptibility. Milled rice varieties with high phenolic content and hardness may show resistance to R.dominica infestation.
However, caution is needed with regard to the introduction of resistant varieties as a method of control of R. dominica; the insect may overcome host plant resistance, as it has developed resistance to insecticides, and the development of further resistance management strategies would be required.
Over the last 15 years, due to environmental concerns and insect pest resistance to conventional chemicals, interest in botanical insecticides has increased. Botanical insecticides are naturally occurring insecticides which are derived from plants (Golob et al., 1999; Isman, 2000). Compared to synthetic compounds they are less harmful to the environment, generally less expensive, and easily processed and used by farmers and small industries.
Botanical insecticides are used in several forms, such as powders, solvent extracts, essential oils and whole plants, these preparations have been investigated for their insecticidal activity including their action as repellents, anti-feedants and insect growth regulators (Weaver and Subramanyam, 2000).
The introduction of powdered leaves of Salvia officinalis L. and Artemisia absinthium L. to wheat grains was very effective in reducing population size and delaying development time of R. dominica (Klys, 2004).
Natural feeding inhibitors found in either wild or cultivated plants are usually alkaloids and glycosides. The mode of action of these compounds is complex and poorly understood, although it is found that insects exposed to such substances usually stop feeding, resulting in a decreased body weight or even death if the insects fail to feed for a long period of time.
Plant essential oils and solvent extracts are the most studied botanical methods of controlling stored grain insect infestations (Stoll, 2000; Shaaya et al., 2003; Moreira et al., 2007; Rozman et al., 2007; Rajendran and Sriranjini, 2008). The essential oils obtained from different plant species repel several insect pests and possess ovicidal and larvicidal properties. Although they are considered by some as environmentally compatible pesticides (Cetin et al., 2004), some botanicals, especially essential oils, are toxic to a broad range of animals, including mammals (Bakkali et al., 2008). Suthisut et al. (2011a,b) found that some natural products were actually more dangerous to use than the commercial insecticides, because much more of the product is needed to control the insects than the fumigant or the synthetic contact insecticide.
Moreira et al. (2007) reported that R. dominica was more susceptible than Sitophilus zeamais and Oryzaephilus surinamensis to hexane crude extract of Ageratum conyzoides, experiencing more than 88% mortality after 24 h of exposure.
Plant oils are also used for their fumigant activity against R. dominica (Lee et al., 2004) on the basis of their efficacy, economic value and use in large-scale storages.
In spite of the wide-spread recognition of insecticidal properties of plants, few commercial products obtained from plants are in use and botanicals used as insecticides presently constitute only 1% of the world insecticide market (Rozman et al., 2007).
The use of natural enemies to control R. dominica and other stored grain insects has been limited in developed countries because of the low tolerance (0-2 insects/kg grain) of insects in stored grain. However, because of the interest in controlling insect pests without the use of insecticides, there has been renewed interest in predators and parasites (Brower et al., 1991). Despite this, research on the potential use of bio-control agents of stored grain insects has been limited to a small number of species.
There have been several laboratory studies on the use of predators of R. dominica (Brower et al., 1991). Teretriosoma nigrescens is a histerid beetle that is found in Central America, where it primarily feeds on Prostephanus truncates, a species closely related to R. dominica. It is able to feed on R. dominica. However, the ability of T. nigrescens to significantly reduce R. dominica populations has yet to be determined (Markham et al., 1994).
Xylocoris flavipes (Hemiptera: Anthocoridae) is a predator of many stored product insect pest (Rahman et al., 2009). The cadelle Tenebroides mauritanicus also feeds on grain, mites and stored-product insect eggs, including Rhyzopertha (Bousquet, 1990). The predatory mites Cheyletus eruditus and Pyemotes ventricosus feed on a wide variety of stored product insect eggs (Asanov, 1980; Brower et al., 1991), but their effect on populations in the field has not been determined. Among the four Cheyletus species found in storage structures of Central Europe, only C. eruditus is employed for the biocontrol of stored grain insect pests (Lukáš et al., 2007).
Parasites and parasitoids
Most of the parasitoids that attack the primary beetle pests are in the families Pteromalidae and Bethylidae. These hymenopteran parasitoids are very small, do not feed on the grain and can easily be removed from the grains by using normal cleaning processes. Choetospila elegans is a small pteromalid wasp that attacks R. dominica and certain other coleopteran and lepidopeteran insect pests. The wasp normally parasitizes larvae that are feeding inside the grain. At 32°C, a wasp takes approximately 15 days to complete its development on R. dominica; the generation time of C. elegans is almost half that of R. dominica. In the presence of hosts, female wasps live for 10-20 days at 32°C. A single female C. elegans is capable of parasitizing up to six R. dominica per day.
During a field study, Flinn et al. (1998) observed that C. elegans suppressed R. dominica by 91% in 27 ton bins of stored wheat compared with control bins. In another study it was found that suppression of R. dominica population growth by parasitoid wasps was significantly higher at 25°C than at 32°C and that 25°C resulted in a very high level of population suppression (99%) compared to the control (Flinn, 1998). Another hymenopteran parasitoid, Anisopteromalus calandrae, is effective at reducing R. dominica populations.
The hymenopteran parasitoid Anisopteromalus calandrae suppressed R. dominica populations in all types of storage bag except those made of polythene. The highest percentage (81%) suppression occurred in calico bags and the lowest suppression (57%) occurred in polypropylene bags (Mahal et al., 2005).
The use of entomopathogenic fungi has been evaluated extensively in laboratory and field studies against R. dominica. The pathogenicity of entomophaghous fungi depends upon various physical (temperature, relative humidity, application time of fungal insecticide, dark and light period etc.) and biological factors (the specific host species, host pathogen interaction etc.). Unlike other microbial control agents, fungi possess the ability to infect the insects through cuticle (Boucias and Pendland, 1991; Thomas and Read, 2007). Beauveria bassiana (Ascomycota: Hyphomycetes) and Metarhizium anisopliae (Ascomycota: Sordario) are the most extensively studies fungal species in this regard (Lord, 2005; Vassilakos et al., 2006; Athanassiou et al., 2008; Wakil and Ghazanfar, 2010).
More recently, various native entomopathogenic fungi, isolated from different components of the maize agroecosystem, how shown virulence against R. dominica and two other stored maize insect species. Paecilomyces and Metarhizium were the most abundant genera isolated from the soil, wheras the isolates of Purpureocillium lilacinum were the best in controlling target insect species (Barra et al., 2013).
Entomopathogenic fungi have also been tested in combination with other control tactics: for example, Isaria fumosorosea with enhanced diatomaceous earth and the plant extract bitterbarkomycin (Riasat et al., 2013); B. bassiana and enhanced diatomaceous earth (Wakil et al., 2011); B. bassiana admixed with a diatomaceous earth formulation (Riasat et al., 2011); and B. bassiana with thiamethoxam and a diatomaceous earth formulation (Wakil et al., 2012). The results demonstrated that such combined controls could be an effective strategy to control R. dominica in stored wheat.
Bacillus thuringiensis var. tenebrionis has been investigated for the control of R. dominica (Keever, 1994; Mummigatti et al., 1994). Most B. thuringiensis varieties are ineffective against beetles; however, R. dominica was one of the more susceptible beetles to B. thuringiensis var. tenebrionis, with more than 75% mortality in 17 days at 250 ppm (Mummigatti et al., 1994). Toxins of 36 available subspecies of B. thuriengensis were evaluated against larvae and adults of R. dominica. The spore crystal complex of B. thuringiensisdarmstadiensis obtained from Germany was the most effective against larvae, but the same subspecies from USA and Japan could not effectively control R. dominica (Beegle, 1996).
In a recent study, the combination of Cry3Aa protoxin and protease inhibitor (potato carboxypeptidase) resulted in delayed development, increased mortality and progeny suppression of R. dominica (Oppert et al., 2011).
Entomopathogenic nematodes (EPNs) are endoparasites of insects (Gaugler, 2002), that enter into host through natural body openings and release mutualistic bacteria inside the host’s body that kills it within 24-48 hours. Their low toxicity to vertebrates (Boemare et al., 1996), exemption from registration in the USA by the Environmental Protection Agency (Kaya and Gaugler, 1993), commercial availability (Grewal, 2002) and ability to seek their host actively (Campbell and Lewis, 2002) make EPNs potentially good biological control agents for stored-product pests. However, they have not proved very effective against R. dominica; R. dominica suffered only 35% adult mortality to two EPN species (in Heterorhabditidae and Steinernematidae) (Ramos-Rodríguez et al., 2006). Similarly, at 200C, the mortality of adult R. dominica in wheat treated with Steinernema feltiae and Steinernema carpocapsae (at 20,000 infected juveniles per ml) did not exceed 23 and 42%, respectively (Athanassiou et al., 2010a).
The insecticidal efficacy of different group of insecticides varies with the surfaces on which they are applied, as insecticides degrade faster on concrete than on wood or metal (Collins et al., 2000). Deltamethrin was more effective on plywood than on concrete or tile surface against R. dominica and Tribolium spp. released for 21 weeks (Arthur, 1997).
Chlorpyrifos-methyl and pirimiphos-methyl, although effective against most stored grain insect pests, are relatively ineffective against R. dominica.
R. dominica was the most susceptible species among the four stored grain insects (R. dominica, T. castaneum, Sitophilus oryzae and Lepinotus reticulatus) to spinosad applied on wheat or maize (Athanassiou et al., 2010b). These findings coincide with that of Vayias et al. (2009), who stated that R. dominica mortality were extremely high on wheat, corn, barley and rice, even when treated low levels of spinosad.
Insect growth regulators (IGR) have low toxicity to mammals, but take longer to control insect populations and are more expensive than other insecticides. They can be sprayed or dusted directly onto the grain, and protect grain from infestation from two weeks to over a year. Methoprene is an IGR commercial formulation labelled in the USA for direct use on to stored grains. In addition to externally feeding stored-grain insects, it is also an effective grain protectant against R. dominica (Arthur, 2004). The combination of methoprene and DE not only eliminated progeny production of R. dominica on stored rice but it also provided a measure of adult control (Chanbang et al., 2007). Kavallieratos et al. (2012) found that IGRs, including two juvenile hormone analogues, four chitin synthesis inhibitors, one ecdysteroid agonist and one combination of chitin synthesis inhibitors and juvenile hormone analogues, tested against R. dominica in wheat resulted in >88.5% suppression of progeny. The highest level of suppression was at 5 ppm. A juvenile hormone analogue (pyriproxyfen) and chitin synthesis inhibitor (lufenuron) suppressed progeny production 100% when applied at 1 ppm, and the highest values of parental mortality were observed in wheat treated with combination of chitin synthesis inhibitors and juvenile hormone analogues (lufenuron + fenoxycarb).
Neonicotinoids are broad spectrum insecticides that can be applied in different ways, such as foliar, soil drench, seed treatment and stem applications, and to various crops (Schulz et al., 2009; Jeschke and Nauen, 2010). Imidacloprid and indoxacarb can control R. dominica, but these chemicals are not yet registered for use on stored grains (Daglish and Nayak, 2012).
The integration of different control measures has been suggested to be one of the most promising approaches to insect management. Daglish and Wallbank (2005) used methoprene + diflubenzeron on stored sorghum and found 98.5-100% reduction in the progeny production of R. dominica four weeks after exposure. On the other hand, the combination of spinosad + s-methoprene could not control methoprene-resistant strain of R. dominica (Daglish, 2008). A combination of thiamethoxam and DE also exhibited promising insecticidal potential against R. dominica in laboratory studies (Wakil et al., 2012).
Phosphine is commonly used as a fumigant, is used to control insect infestations in stored commodities. Although it is effective, commodities can become re-infested once the fumigant has dissipated. Phosphine is also highly toxic to humans and should be handled with extreme care. It is usually applied to the grain as aluminium phosphide pellets or tablets, although magnesium phosphide is also available in some countries. Some countries allow phosphine to be delivered to the grain from compressed gas cylinders. A system has been developed in Australia in which phosphine mixed with carbon dioxide is delivered to the grain at low concentrations and continuous flow for several weeks. At high temperatures and humidity regimes, phosphine is corrosive to copper and can cause damage to electrical systems.
After treatment with an insecticide, grain often must be held for a certain period of time before it can be processed or used as animal feed. The period of protection is dependant upon the commodity treated, the temperature, grain moisture content and the insecticide used. In general, temperatures must be over 15°C for effective control; higher temperatures cause more rapid control but also cause faster degradation of the insecticide. High moisture content also reduces the duration of protection. Many of these insecticides can be used as a structural treatment to eliminate residual insect populations from empty silos and buildings.
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21/08/14 Text updated by:
Waqas Wakil, University of Agriculture, Faisalabad, Pakistan
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