Prostephanus truncatus (larger grain borer)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- 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
- Means of Movement and Dispersal
- Pathway Vectors
- Plant Trade
- Impact Summary
- Environmental Impact
- Impact: Biodiversity
- Social Impact
- Detection and Inspection
- Prevention and Control
- Links to Websites
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Prostephanus truncatus (Horn)
Preferred Common Name
- larger grain borer
Other Scientific Names
- Dinoderus truncatus Horn
International Common Names
- English: grain, borer, larger; greater grain borer; scania beetle
- Spanish: barrebador de los granos; barrenador del grano mayor; barrenador grande de los graneros
Local Common Names
- Germany: Bohrer, grosser Korn-
- PROETR (Prostephanus truncatus)
Summary of InvasivenessTop of page
On a local scale, the pest flies from environments where population density is high or food exhausted to seek new hosts, usually in subsistence farmers' granaries. It locates stored product hosts, such as maize and dried cassava either by chance or because they are already infested and harbour males releasing aggregation pheromone. This type of host selection leads to very dense populations of the pest in a few farm stores surrounded by other stores showing no signs of infestation.
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Arthropoda
- Subphylum: Uniramia
- Class: Insecta
- Order: Coleoptera
- Family: Bostrichidae
- Genus: Prostephanus
- Species: Prostephanus truncatus
Notes on Taxonomy and NomenclatureTop of page
The most commonly used English name for the species is larger grain borer (LGB) although some countries favour greater grain borer (GGB) which gives the correct semantic distinction from the lesser grain borer (Rhyzopertha dominica).
The adults may be identified using the keys of Fisher (1950), Kingsolver (1971) or Haines (1991). A key to both larvae and adults is given in Gorham (1991). The taxonomy, systematics and identification of P. truncatus have been reviewed recently by Farrell and Haines (2002).
DescriptionTop of page
The larvae are white, fleshy and sparsely covered with hairs. They are parallel-sided, i.e. they do not taper. The legs are short and the head capsule is small relative to the size of the body.
The adult has the typical cylindrical bostrichid shape. The declivity is flattened and steep and has many small tubercles over its surface. The limits of the declivity, apically and laterally, are marked by a carina. The antennae are 10-segmented and have a loose three-segmented club; the 'stem' of the antenna is slender and clothed with long hairs and the apical club segment is as wide as, or wider than, the preceding segments. The body is 3-4.5 mm long.
DistributionTop of page
P. truncatus is indigenous in Central America, tropical South America, and the extreme south of the USA as a major, but localized, pest of farm-stored maize. It was introduced into Tanzania, probably in the late 1970s, and has become a serious pest of stored maize and dried cassava in that part of East Africa; it has since spread into Kenya, Burundi, Rwanda, Malawi, Zambia, Mozambique, Namibia and South Africa, and is almost certainly present but unreported from several other countries in the region. It was first found in West Africa in Togo in 1984 and it has since spread to Benin, Nigeria, Ghana, Niger and Burkina Faso. A separate outbreak occurred in Guinea Conakry.
P. truncatus could potentially invade all maize and cassava growing areas of tropical and sub-tropical Africa, and it is the only recent example of a serious storage pest invading on a regional or continental scale. It remains a quarantine threat to other maize growing regions in the Old World.
There are several records of interceptions of P. truncatus, including Canada (Manitoba), the USA (Arizona, Montana, New York and New Jersey), Germany (interception from Guatemala), Israel and Iraq (IIE, 1995).
A record of P. truncatus in Thailand is now known to be based on a misidentification (R. Hodges, Natural Resources Institute, UK, personal communication, 2006).
A record of P. truncatus in Peru (Hodges, 1986) published in previous versions of the Compendium is invalid as it is based on personal correspondence cited in Wright (1984). SENASA has developed a specific surveillance programme for grain-stored pests in Peru. P. truncatus has not been detected in any surveys conducted by SENASA since the surveillance programme began in 1999 (SENASA, 2021).
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: 13 May 2022
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Burkina Faso||Present, Localized||Introduced||1991|
|South Africa||Present, Localized|
|Tanzania||Present, Widespread||Introduced||Invasive||First reported: <1981|
|China||Absent, Intercepted only|
|Hong Kong||Absent, Intercepted only|
|India||Absent, Intercepted only|
|-Uttar Pradesh||Absent, Intercepted only|
|Iraq||Absent, Intercepted only|
|Israel||Absent, Intercepted only|
|Philippines||Absent, Intercepted only|
|Thailand||Absent, Invalid presence record(s)|
|France||Absent, Intercepted only|
|Germany||Absent, Intercepted only|
|Canada||Absent, Intercepted only|
|-Manitoba||Absent, Intercepted only|
|United States||Present, Localized|
|-District of Columbia||Present||Introduced||1878|
|-Montana||Absent, Intercepted only|
|-New Jersey||Absent, Intercepted only|
|-New York||Absent, Intercepted only|
|Brazil||Absent, Formerly present||1993|
|-Minas Gerais||Absent, Formerly present|
|Peru||Absent, Invalid presence record(s)|
History of Introduction and SpreadTop of page
Risk of IntroductionTop of page
Habitat ListTop of page
Hosts/Species AffectedTop of page
P. truncatus behaves as a typical primary pest of farm-stored maize; whole grains are attacked, on the cob, both before and after harvest. P. truncatus is also a pest of farm-stored cassava, particularly cassava chips. The adults bore into a wide range of foodstuffs and other materials including wood, bamboo, plastic and soap. Extensive populations of P. truncatus occur in the natural environment, and it has been recorded from a number of tree species in Central America (Rees et al., 1990; Ramirez-Martinez et al., 1994) and Africa (Nang'ayo et al., 1993, 2002; Nansen et al., 2004) in some cases associated with twig girdling by cerambycid beetles (Borgemeister et al., 1998).
Host Plants and Other Plants AffectedTop of page
|Arachis hypogaea (groundnut)||Fabaceae||Other|
|Manihot esculenta (cassava)||Euphorbiaceae||Main|
|Phaseolus vulgaris (common bean)||Fabaceae||Other|
|Sorghum bicolor (sorghum)||Poaceae||Other|
|stored products (dried stored products)||Main|
|Triticum aestivum (wheat)||Poaceae||Other|
|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
Adults frequently initiate their attack on stored maize cobs with intact sheaths by boring into the base of the maize cob cores, although they eventually gain access to the grain via the apex of the cob by crawling between the sheathing leaves (Hodges and Meik, 1984). Adults bore into the maize grains, making neat round holes, and as they tunnel from grain to grain they generate large quantities of maize dust. Adult females lay eggs in chambers bored at right angles to the main tunnels. Egg-laying on stabilized grain, like that on the maize cob, is more productive than on loose-shelled grain as the oviposition period is longer, equal in length to the life of the female, and the eggs are laid at a greater rate.
Larvae hatch from the eggs after about three days at 27°C and seem to thrive on the dust produced by boring adults. For example, large numbers of larvae develop and pupate in dust at the base of dense laboratory cultures.
The life cycle of P. truncatus has been investigated at a range of temperatures and humidities (Shires, 1979, 1980; Bell and Watters, 1982; Hodges and Meik, 1984). Development of the larva through to the adult stage at the optimum conditions of 32°C and 80% RH takes 27 days on a diet of maize grain. Humidity within the range 50-80% RH does not greatly affect the development period or mortality; at 32°C, a drop in RH from 80 to 50% (giving maize with an equilibrium moisture content of about 10.5%) extended the mean development period by just 6 days and increased the mean mortality by only 13.3%. This tolerance of dry conditions was confirmed during field studies in Nicaragua and Tanzania in which maize at 10.6 and 9% moisture content, respectively, was heavily infested.
The success of this pest may be partly due to its ability to develop in grain at low moisture. Many other storage pests are unable to increase in number under low moisture conditions. For example, Sitophilus oryzae, a species occurring in the same ecological niche, needs a grain moisture content of at least 10.5% for development. Thus, in dry conditions, P. truncatus probably benefits from the absence of any significant competition from other storage pests.
P. truncatus develops more rapidly on maize grain than on cassava; at 27°C and 70% RH, the respective development periods on maize grain and cassava were 32.5 and 40 days, respectively. Under ideal conditions of temperature and humidity on maize cobs or stabilized maize grain, estimates for the intrinsic rate of increase (r) of P. truncatus are in the order of 0.7-0.8 per week; this is similar to the rate of increase reported for Tribolium castaneum under comparable climatic conditions.
Details of flight performance and factors affecting flight and distribution behaviour have been investigated in the laboratory (Fadamiro and Wyatt, 1955, 1996; Fadamiro, 1997). A field study in Honduras showed flight activity of P. truncatus following a daily bimodal pattern with a major peak at 06.00-08.00 h and a minor peak at 18.00-20.00 h (Novillo, 1991). A similar pattern was observed by Tigar et al. (1993) in Central Mexico and Birkinshaw et al. (2004) in Ghana, but in both these cases the major peak was associated with dusk.
Adults may be sexed using a method described by Shires and McCarthy (1976).
For further detailed information on biology and ecology, consult reviews by Hodges (1986), Markham et al. (1991), Hodges (1994), Nansen and Meikle (2002) and Hill et al. (2002).
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
|Teretrius nigrescens||Predator||Eggs; Arthropods|Larvae; Arthropods|Pupae||Africa; Togo||stored maize|
Notes on Natural EnemiesTop of page
Infestations of P. truncatus in Nicaragua were associated with high levels of parasitic Hymenoptera, accounting for 10% of the total number of insects present, while the P. truncatus population represented 56.5%; these Hymenoptera were not identified. In Tanzania, large numbers of Anisopteromalus calandrae were associated with P. truncatus when few other potential hosts were present (Hodges et al., 1983). The hemipteran Xylocoris flavipes has been observed as a predator of all three larval stages of P. truncatus in West Africa, however populations of the bug declined as pest numbers rose. It is believed that the conditions created by P. truncatus infestation are unfavourable to X. flavipes and hence this species probably does not play an important role in the control of the pest (Helbig, 1999).
Little is known about the relationship between P. truncatus and other organisms. Infestations of this beetle are found, together with those of other species, but P. truncatus is the predominant storage species in the dry conditions of Tanzania and Nicaragua. It has been demonstrated that P. truncatus is deterred from infesting grain in which the larvae of Sitophilus zeamais are developing and that this probably results from substances deposited on the grain surface by adult S. zeamais (Danho et al., 2000).
In a survey of the pest in Kenya, Odour et al. (2000) found that the entomopathogenic fungus Beauveria bassiana occurred on only 0.08 to 0.94% of the total insects collected. This is a low infection rate. Isolates of Beauveria, Metarhizium and Paecilomyces obtained in Ethiopia where all found to be virulent against P. truncatus although a total immersion bioassay was used so this result might have been anticipated (Kassa et al., 2002). The potential of Beauveria has been investigated in field trials designed to give a better understanding of how future research efforts can develop methods to make this fungus an effective means of control for P. truncatus (Meikle et al., 2001).
Means of Movement and DispersalTop of page
Pathway VectorsTop of page
|Land vehicles||Carried by national and international grain trade||Yes|
Plant TradeTop of page
|Plant parts liable to carry the pest in trade/transport||Pest stages||Borne internally||Borne externally||Visibility of pest or symptoms|
|Roots||arthropods/adults; arthropods/eggs; arthropods/larvae; arthropods/pupae||Yes||Pest or symptoms usually visible to the naked eye|
|True seeds (inc. grain)||arthropods/adults; arthropods/eggs; arthropods/larvae; arthropods/pupae||Yes||Pest or symptoms usually visible to the naked eye|
|Wood||arthropods/adults; arthropods/eggs; arthropods/larvae; arthropods/pupae||Yes||Pest or symptoms usually visible to the naked eye|
|Plant parts not known to carry the pest in trade/transport|
|Fruits (inc. pods)|
|Growing medium accompanying plants|
|Stems (above ground)/Shoots/Trunks/Branches|
Impact SummaryTop of page
|Fisheries / aquaculture||None|
ImpactTop of page
Infestations in maize may start on the mature crop in the field, i.e. when moisture content is at or below 18%. Weight losses of up to 40% have been recorded in Nicaragua from maize cobs stored on the farm for 6 months (Giles and Leon, 1975). In Tanzania, up to 34% losses have been observed after 3 months storage on the farm, with an average loss of 8.7% (Hodges et al., 1983). P. truncatus is a much more damaging pest when compared to other storage insects including Sitophilus oryzae, S. zeamais and Sitotroga cerealella, under similar conditions; maize losses due to these other species were 2-6, 3-5 and 2-5%, during a storage season in Zambia, Kenya and Malawi, respectively.
Losses caused by P. truncatus in dried cassava roots can be very high; the dried roots are readily reduced to dust by boring adults and a loss of 70% has been recorded after only 4 months of farm storage (Hodges et al., 1985). A group of 25 farmers from five villages in Togo sustained average cumulative losses of 9.7% after 3 months storage, this figure rose to 19.5% after 7 months (Wright et al., 1993).
Not all problems with this pest are restricted to farmers' granaries. In the early days after the arrival of P. truncatus in East Africa, countries with the pest found their maize exports banned. For example in 1987-88, it is estimated that Tanzania lost US$634,000 in export earning. This situation improved following efforts to upgrade phytosanitary procedures in the region but such procedures, involving fumigation, have their own continuing costs (Boxall, 2002).
A grain injury model for P. truncatus infesting farm-stored maize in West Africa has been developed at the International Institute of Tropical Agriculture. It can be used in conjunction with predictive models of pest population dynamics to guide the development of integrated pest management strategies (Holst et al., 2000a). The models are conveniently displayed, together with information on sampling routines, on a web site (<_http3a_ _www.agrsci.dk2f_plb2f_bembi2f_africa2f_project.htm="" xmlns="http://cabi.org/cet">).
A detailed review of the damage and loss caused by P. truncatus has been prepared by Boxall (2002). He considers loss of value, nutrition as well as impact at the national level, effect on international trade and modern methods of rapid loss assessment. In the early days after the arrival of P. truncatus in East Africa, countries with the pest found their maize exports banned. For example, in 1987-1988, it is estimated that Tanzania lost US$634,000 in export earning. This situation improved following efforts to upgrade phytosanitary procedures in the region but such procedures, involving fumigation, have their own continuing costs.
Environmental ImpactTop of page
Impact: BiodiversityTop of page
Social ImpactTop of page
Detection and InspectionTop of page
Flight traps, such as funnel, delta or wing traps baited with the male-released aggregation pheromone of P. truncatus, are highly effective for monitoring this species. These traps should be placed at least 100 m from stores, which contain maize or dried cassava, or from the standing maize crop to avoid attracting the beetles to these food sources. A detailed leaflet giving recommendations on the use of pheromone traps to monitor P. truncatus has been prepared by Hodges and Pike (1995). For long-term, routine trapping programmes in both East and West Africa the Japanese beetle flight-trap baited with P. truncatus pheromone is now the method of choice while pheromone baited Delta traps, made from cardboard, are used in short-term programmes.
The risk of stores becoming infested by P. truncatus is known to be related to the number of P. truncatus that are flying (Birkinshaw et al., 2002). There are big variations both within and between years in the numbers of beetles taking flight. In many locations this difference between years is noticeable by the extent to which farmer's stores become infested, i.e. there are good and bad years. A computer-based model has been developed that uses climate data to predict P. truncatus flight activity and in this way it can predict the years when P. truncatus infestation will be bad (Hodges et al., 2003).
Monitoring for the presence of P. truncatus in farm maize stores themselves is difficult because the pest is not attracted to its pheromone when present on its food. A sequential sampling plan for inspecting stores in West Africa has been described by Meikle et al. (2000). Compton and Sherington (1999) have described a rapid method for estimating the weight losses caused by P. truncatus to maize cobs.
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.
The most effective method of controlling P. truncatus in maize is to admix a dilute dust insecticide. P. truncatus is highly susceptible to synthetic pyrethroid insecticides such as permethrin and deltamethrin. However, these insecticides are relatively ineffective against other storage pests such as Sitophilus spp. and Tribolium castaneum, which occur in the same pest complex and are more susceptible to organophosphorus insecticides. Both types of insecticide are applied in order to control the whole complex (Golob et al., 1985; Golob and Hanks, 1990). Combinations such as pirimiphos-methyl and permethrin, deltamethrin and pirimiphos-methyl or fenitrothion and fenvalerate have been used successfully to protect farm-stored grain. Fumigation with phosphine is very effective in large-scale stores.
Recent laboratory and field studies have shown that unless inert dusts are applied at very high rates, they are not particularly effective against P. truncatus. However, good control can be achieved when they are mixed with insecticides or soil bacteria metabolites such as Spindeba (Stathers, 2003).
For a detailed review of chemical, physical and cultural methods for the control of P. truncatus, see Golob (2002).
Detailed reviews on the control of P. trunctus by the predator Teretrius nigrescens have been published by Meikle et al. (2002) and Borgemeister et al. (2003). Initial releases of T. nigrescens were in Togo in 1991 and in Kenya in 1992. In both countries it became well established and spread. Subsequently, there have been predator releases in Benin, Ghana, Tanzania and Malawi. Only in the case of Tanzania does it appear that there has been any difficulty in the predator becoming quickly and easily established. However, despite the successful introductions, there are still regular outbreaks of P. truncatus and farmers still suffer losses. It has been concluded by Holst et al. (2000b) that T. nigrescens does not offer a good example of classical biological control but as the predator is able to reduce the density of the pest it is considered that it has, nevertheless, a role to play in integrated pest management.
Cultural Control and Sanitary Methods
Good store hygiene, especially the removal of infested residues and the selection of only sound material for storage, can play an important role in limiting infestation by P. truncatus. The use of resistant cultivars may also reduce the severity of an infestation, although much work remains to be done on the mechanisms of resistance.
ReferencesTop of page
Adda C, Borgemeister C, Meikle WG, Markham RH, Olaleye I, Abdou KS, Zakari MO, 1996. First record of the larger grain borer, Prostephanus truncatus (Coleoptera: Bostrichidae), in the Republic of Niger. Bulletin of Entomological Research, 86(1):83-85; 11 ref
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Back EA, Cotton RT, 1938. Stored Grain Pests. Farmers' Bulletin No. 1260 (revised). Washington DC, USA: US. Department of Agriculture
Bell RJ, Watters FL, 1982. Environmental factors influencing the development and rate of increase of Prostephanus truncatus (Horn) (Coleoptera: Bostrichidae) on stored maize. Journal of Stored Products Research, 18(3):131-142
Birkinshaw LA, Hodges RJ, Addo S, 2004. Flight behaviour of Prostepahanus truncatus and Teretrius nigrescens demonstrated by a cheap and simple pheromone-baited trap designed to segregate catches with time. Journal of Stored Products Research, 40:227-232
Borgemeister C, Goergen G, Tchabi A, Awande S, Markham RH, Scholz D, 1998. Exploitation of a woody host plant and cerambycid-associated volatiles as host-finding cues by the larger grain borer (Coleoptera: Bostrichidae). Annals of the Entomological Society of America, 91(5):741-747; 44 ref
Borgemeister C, Holst N, Hodges RJ, 2003. Biological control and other pest management options for larger grain borer Prostephanus truncatus. In: Neuenschwander P, Boregemeister C, Langewald J, eds. Biological Control in IPM Systems in Africa. Wallingford, UK: CABI Publishing, 311-328
Boxall RA, 2002. Damage and loss caused by the Larger Grain Borer, Prostephanus truncatus (Horn) (Coleoptera: Bostrichidae. Integrated Pest management Reviews, 7:105-121
Danho M, Haubruge E, Gaspar C, Lognay G, 2000. Selection of grain-hosts by Prostephanus truncatus (Coleoptera, Bostrychidae) in the presence of Sitophilus zeamais (Coleoptera, Curculionidae) previously infested grains. Belgian Journal of Zoology, 130(1):3-9
DCA - Nationalt Center for Fødevarer og Jordbrug, Denmark, 2018. Update of pest status in Denmark for specific harmful organisms in relation to export of seeds - part 1. (Opdatering af skadegørerstatus i Danmark for specifikke skadegørere i relation til eksport af frø - del 1). Tjele, Denmark: DCA - Nationalt Center for Fødevarer og Jordbrug, Aarhus University.51 pp. https://pure.au.dk/portal/files/141699745/Levering_af_skadeg_rerestatus_del_1_ver5.pdf
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Hodges RJ, Birkinshaw LA, Smith RH, 1998. Host selection or mate selection? Lessons from Prostephanus truncatus, a pest poorly adapted to stored products. Proceedings of the 7th International Working Conference on Stored products Protection. Beijing China, October 1998, in press
Hodges RJ, Farrell G, Golob P, 1996. Review of the Larger Grain Borer outbreak in East Africa - rate of spread and pest status. In: Farrell G, Greathead AH, Hill MG, Kibata GN, eds. Management of Farm Storage Pests in East and Central Africa. Proceedings of the East and Central Africa Storage Pest Management Workshop, Naivasha, Kenya, 14-19 April 1996. Egham, UK: International Institute of Biological Control, 29-43
Hodges RJ, Meik J, Denton H, 1985. Infestation of dried cassava (Manihot esculenta Crantz) by Prostephanus truncatus (Horn) (Coleoptera: Bostrichidae). Journal of Stored Products Research, 21(2):73-77
Hodges RJ, Pike V, 1995. How to use pheromone traps to monitor the Larger Grain Borer (Prostephanus truncatus). Chatham Maritime, Kent, UK: Natural Resources Institute
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Holst N, Meikle WG, Markham RH, 2000. Grain injury models for Prostephanus truncatus (Coleoptera: Bostrichidae) and Sitophilus zeamais (Coleoptera: Curculionidae) in rural maize stores in West Africa. Journal of Economic Entomology, 93(4):1338-1346; 23 ref
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Kassa A, Zimmermann G, Stephan D, Vidal S, 2002. Susceptibility of Sitophilus zeamais (Motsch.) (Coleoptera: Curculionidae) and Prostephanus truncatus (Horn) (Coleoptera: Bostrichidae) to entomopathogenic fungi from Ethiopia. Biocontrol Science and Technology, 12(6):727-736; 21 ref
Kingsolver JM, 1971. A key to the genera and species of Bostrichidae commonly intercepted in USDA plant quarantine inspections. USDA Agricultural Quarantine Inspection Memorandum, No. 697
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