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7 June 2022

Trogoderma granarium (khapra beetle)

Datasheet Types: Pest, Invasive species

Abstract

This datasheet on Trogoderma granarium covers Identity, Overview, Distribution, Dispersal, Hosts/Species Affected, Diagnosis, Biology & Ecology, Environmental Requirements, Seedborne Aspects, Natural Enemies, Impacts, Prevention/Control, Further Information.

Identity

Preferred Scientific Name
Trogoderma granarium Everts
Preferred Common Name
khapra beetle
Other Scientific Names
Trogoderma affrum
Trogoderma khapra Arrow
International Common Names
English
beetle, khapra
Spanish
dermeste de los granos
escarabajo khapra
gorgojo de khapra
gorgojo khapra
French
dermeste des grains
trogoderme
trogoderme des grains
Local Common Names
Germany
Khapra-Käfer
Greece
trogoderma ton sitiron
Israel
chipushit hagargirim
Italy
dermeste dei cereali
Norway
khaprabille
Turkey
kapra

Pictures

Adults oval, 2-3 mm, females somewhat larger than males, dorsal surface moderately clothed in fine hairs.
Adult
Adults oval, 2-3 mm, females somewhat larger than males, dorsal surface moderately clothed in fine hairs.
NRI/MAFF
Adults reddish-brown, with or without vague darker markings, pronotum darker brown. Larvae typically very hairy with a 'brush' of long spicisetae like a tail; ranging from ca 1.6 mm long (first instar) to 5 mm (fully mature).
Adult and larva
Adults reddish-brown, with or without vague darker markings, pronotum darker brown. Larvae typically very hairy with a 'brush' of long spicisetae like a tail; ranging from ca 1.6 mm long (first instar) to 5 mm (fully mature).
©NRI/MAFF
Adult, larva and larval skins of T. granarium, and damage to wheat grains.
Adult and larva on wheat
Adult, larva and larval skins of T. granarium, and damage to wheat grains.
Ministry of Agriculture

Summary of Invasiveness

The khapra beetle, T. granarium, is a serious pest of stored products, and has been classified among the 100 most important invasive stored product species. Under optimum conditions for its development, the population of T. granarium can grow enormously in a very short period of time, and it can easily outcompete other major stored product insect species of stored grains. One of the key elements in its biology is a long diapause at the larval stage, which can be interrupted for ‘foraging excursions’. Moreover, diapausing larvae are generally much more tolerant to certain insecticides and non-chemical treatments, as compared with non-diapausing ones. The identification of T. granarium is difficult, as it is morphologically very similar to other species of the genus Trogoderma. It is considered a quarantine species in many parts of the world, while interceptions during the past few years have notably increased, indicating a serious risk for further spread and the need for increased surveillance.

Taxonomic Tree

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Notes on Taxonomy and Nomenclature

The family Dermestidae contains a number of species that are pests of stored products, the most serious of which is T. granarium. Larval and adult keys are available for the identification of species of economic importance in North America (Gorham, 1987), for use in the tropics (Haines, 1991) and for the identification of adults of the few stored product species occurring in the UK (Green, 1979). It is possible to separate the one important pest species, T. granarium, from others that are likely to occur in the tropics, but confirmation of the identification of this species often requires dissection of the mouth parts or genitalia (Green, 1979). For further information, see Hinton (1945) and Peacock (1993); the latter has both larval and adult keys to all dermestids recorded from the UK as well as general information on their biology.
Considering the difficulties in the identification of this species, recent studies have proposed molecular diagnostics to separate T. granarium from other species of this genus, in both laboratory screenings and field scale surveys (Olson et al., 2014; Castañé et al., 2020).

Description

Larva
The larvae are typically very hairy. Spicisetae of various lengths are arranged over the dorsal surface and a 'brush' of long spicisetae on the ninth abdominal segment projects posteriorly like a tail; the length of this brush decreases relative to body size as the larvae grow. Hastisetae are present, and are inserted on the tergites, often in distinct tufts. The first-instar larva is yellowish white, about 1.6 mm long, and has two tufts of 4-10 hastisetae on each of the seventh and eighth abdominal tergites. On reaching the fourth instar, the larvae become golden brown, measure about 3 mm in length, and have dense tufts of hastisetae inserted on the posterolateral parts of the abdominal and thoracic tergites; the tufts become larger and denser posteriorly. As the larva continue to develop, no further changes of setation occur. When fully mature, the larva measures about 5 mm, but can vary in size, especially in the case of diapausing larvae.
Adult
Adult T. granarium are reddish brown, with or without vague darker markings; the pronotum is a darker brown. They are oval in shape, and vary in size from 2 to 3 mm, the females being somewhat larger than the males. The dorsal surface is moderately clothed in fine hairs. A median ocellus is present between the compound eyes. The number of antennal segments is usually 11, but some fusion of the segments may take place so that there can be as few as nine. The fairly distinct antennal club consists of 3-5 segments, depending on the degree of fusion of the distal segments. In the male, the apical segment of the club is elongated in comparison with that of the female. The antennae fit into ventral grooves in the prothorax.

Distribution

The native distribution of T. granarium is not known for certain, but is believed to be the Indian subcontinent, although some authors dispute this; see Banks (1977) for a review. Its distribution has been reviewed by Banks (1977) and more recently by Athanassiou et al. (2019). The beetle usually occurs in hot, dry conditions, predictably in areas which, for at least 4 months of the year, have a mean temperature greater than 20°C and an RH below 50%. It is especially prevalent in certain areas of the Middle East, Africa and South Asia, and is also found in certain specialized warm habitats in temperate countries, e.g. maltings in the UK, although it may no longer occur in the UK (Peacock, 1993). T. granarium does not appear to be established in South East Asia, South America or Australia. Nevertheless, the latest map from the European and Mediterranean Plant Protection Organization (EPPO) shows that this species is established in Africa, and also in other areas of the Eastern Mediterranean zone, such as Cyprus and Turkey (EPPO, 2013; 2016; 2021).

Distribution Map

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Distribution Table

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History of Introduction and Spread

Hinton (1945) listed its distribution as ‘India, Sri Lanka, Malaya’, with introductions into Europe and Asia, etc. Presumably, therefore, records for Africa and the Middle East are mostly more recent. Nevertheless, in an extensive survey of the presence of stored product beetles in ship holds introduced in the UK, Aitken (1975) underlines the absence of T. granarium. From the data available so far, the species is not established in large geographical areas, such as North America and Australia (Athanassiou et al., 2019).
In the first half of the 20th century, there are several records of introductions of T. granarium in Europe in imports from areas that are now India and Pakistan (Athanassiou et al., 2019 and references therein), and quickly spread in different structures in the UK and Germany (Durrant, 1921; Voelkel, 1924; Zacher, 1938; Munro, 1940; Howe and Freeman, 1955; Burges, 1959). It is estimated that T. granarium had been introduced in the USA, in the state of California, during the 1940s, but was misidentified at first as the black carpet beetle, Attagenus piceus [Attagenus unicolor], and this mistake provided the time that was needed for T. granarium to establish in several other states, until its correct identification, which resulted in a very costly eradication programme, by the United States Department of Agriculture (USDA) (Armitage, 1954; 1958; Beal, 1956; Pasek, 1998). However, after its eradication, the species has been found again in a large geographical area in USA, resulting in another series of eradication actions (Pasek, 1998; Day and White, 2016).
Errors in insect identification and regulatory ambiguities caused the inclusion of Australia in the list of the ‘khapra beetle countries’ (Bailey, 1958; 1965). Based on this, it took several years of efforts and lobbying to remove Australia from this list, which involved a position statement from FAO (Bailey, 1965). Due to their dry climate, large areas of Australia are considered suitable for the establishment of this species (Emery et al., 2008; Day and White, 2016).
In 1980, T. granarium was recorded in rice silos in central areas of Taiwan. The pest was eradicated and has not been reported since 1990 in any nationwide studies of stored grain insects in Taiwan (Yao et al., 2016). In 2012, the Bureau of Animal and Plant Health Inspection and Quarantine (BAPHIQ) conducted a three-year monitoring survey for T. granarium in paddy silos, imported rice bins and rice mills in Taiwan. The results indicated that T. granarium is no longer present in Taiwan (Yao et al., 2016). T. granarium is listed as a quarantine pest in Taiwan.
Athanassiou et al. (2019) provides an updated map of countries where the presence of T. granarium has been detected, including the countries where this species is established, and the countries where interceptions have been recorded. From these data, it is indicated that the species is established in most of Africa, and in a zone from the Middle East to India and Myanmar, and also in Korea Republic and some parts of Europe. However, recent surveillances show that the species might not occur in areas that had been officially indicated that the T. granarium is present. For instance, in a recent survey in Spain, Castañé et al. (2020) found that, despite what was believed, T. granarium was not detected, and most individuals of this genus belonged to the larger cabinet beetle, Trogoderma inclusum. Considering the continuous interceptions of this species in a wide range of geographical areas, and the fact that its establishment patterns have to be revisited, there is a need for updated surveillance efforts to illustrate its actual distribution.

Risk of Introduction

Trogoderma granarium is an A2 pest in the European and Mediterranean Plant Protection Organization (EPPO) region (Smith et al., 1992). This suggests that EPPO prioritizes this pest as a potential threat; however, there are now national surveys in the EU towards this direction, despite the fact that its distribution in Europe is poorly understood (Castañé et al., 2020).
Day and White (2016) provide an extensive review of the interceptions and the eradication efforts in Australia and also in other parts of the world, illustrating the risk for certain areas. The risk of introduction of T. granarium through the international marine shipping network in relation of entry ports has been estimated by Paini and Yemshanov (2012). In that study, the authors highlight the risk of introduction of T. granarium to Australia and certain areas of Asia. Similar risk considerations have been reported for invasion pathways in the south-eastern USA (Pasek, 1998; French and Venette, 2005; Athanassiou et al., 2019).

Means of Movement and Dispersal

Natural Dispersal

Given that the adult of T. granarium does not fly, its dispersal through adults that move among different storage and processing facilities is slow. Most dispersal is due to the movement of larvae, through international trade of different commodities and materials, and also through personal items (Athanassiou et al., 2019). The fact that its dispersal is relatively slow, can be an advantage in eradication programmes.

Accidental Introduction

There are certain paradigms of accidental introduction of T. granarium in the USA, which resulted in an expensive but successful eradication programme (Athanassiou et al., 2019). Moreover, another species of Dermestidae has been misidentified as T. granarium in Australia, causing a time-consuming procedure to confirm that eventually this country was khapra beetle-free. During the last decade, however, the number of interceptions of this species has increased in certain areas (Athanassiou et al., 2019).

Plant Trade

Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
True seeds (inc. grain)
Arthropods/Larvae
 YesPest or symptoms usually visible to the naked eye
Plant parts not known to carry the pest in trade/transport
True seeds (inc. grain)

Hosts/Species Affected

The larvae of T. granarium are serious pests of oilseeds, damaged cereals and, to a lesser extent, pulses. The adults rarely, if ever, eat or drink. Females can produce eggs without having fed once emerged from pupae (Hinton, 1945).
Adults that have emerged from diapausing larvae can produce a larger number of eggs than adults emerged from non-diapausing larvae, contributing to a faster population growth right after the termination of the diapause (Karnavar, 1972). T. granarium can easily outcompete other major stored product insect species on grains, such as the lesser grain borer, Rhyzopertha dominica, and the rice weevil, Sitophilus oryzae, at temperatures that are 30ºC or higher (Kavallieratos et al., 2017a). At these temperatures, on wheat, few parental adults of this species can produce thousands of offspring, turning the kernels into frass, in only 65 days (Kavallieratos et al., 2017a). Like many other Dermestidae of stored products, larvae of T. granarium are scavengers at late stages of the ecological succession of storage ecosystems, and thus, they are considered as ‘dirty feeders’. Still, T. granarium larvae can be primary colonizers, damaging more grain than they actually consume. Larvae of this species can consume cast skins, etc. but cannot feed on flour made by the yellow mealworm, Tenebrio molitor, in contrast with other stored product insects that can feed on T. molitor flour (Rumbos et al., 2020).

Host Plants and Other Plants Affected

Growth Stages

Post-harvest

Symptoms

Trogoderma granarium may remain hidden deep in the stored food for relatively long periods. In bag stores, the first signs of infestation are masses of hairy cast larval skins, which gradually push out from the crevices between sacks; this is a sign that the stored food should be fumigated immediately. The larvae crawl over and consume the grain.
Cast skins indicate the presence of larvae of T. granarium, but this happens in the case of several species of this genus that infest stored products (Athanassiou et al., 2019). The accurate identification of T. granarium is essential, as the infestation patterns in stored products are similar with those of other species, such as the warehouse beetle, Trogoderma variabile.

List of Symptoms/Signs

Symptom or signLife stagesSign or diagnosisDisease stage
Plants/Seeds/external feeding   

Diagnosis

The identification of T. granarium is a demanding procedure and has to be performed by trained personnel (Athanassiou et al., 2019). At the adult stage, T. granarium can be confused with other relative species, such as Trogoderma glabrum or Trogoderma variabile (Athanassiou et al., 2019). Morphological identification at the larval stage is even more complicated, as it is based on characters that are not easily recognizable, such as hastisetae.
Certain molecular diagnostics have been proved accurate in separating T. granarium individuals from other species of the genus Trogoderma, and have been used with good results in field sampling (Castañé et al., 2020).

Similarities to Other Species/Conditions

As indicated above, both adults and larvae of T. granarium can be misidentified as other species of this genus, with might cause a ‘false’ interception alarm. These species can be accurately identified by trained personnel under the stereoscope, or through molecular markers (Olson et al., 2014). Correct identification consists of a reliable monitoring system, and provide the inferences necessary for effective eradication strategies. In this context, the increase in the numbers of interceptions of T. granarium might be overestimated, as some specimens that are identified as T. granarium may eventually belong to other species of this genus.

Habitat

Hagstrum and Subramanyam (2009) provide a list of commodities and substances that have been found to be infested by T. granarium, a list that was later completed by Athanassiou et al. (2016) and Kavallieratos et al. (2019). Based on these data, T. granarium can infest more than 100 different commodities of both plant and animal origin, ranging from grains to milk powder. Regarding cereal grains and pulses, T. granarium can develop much easier in wheat than on other products, such as maize, rice or legumes (Athanassiou et al., 2016; Kavallieratos et al., 2019), and thus, the species can be considered as a major threat for cereals and related amylaceous products (Athanassiou et al., 2019).
Development of T. granarium was better on cracked wheat and also in flour, as compared with herbs, pulses, tobacco and dried fruit (Athanassiou et al., 2016; Kavallieratos et al., 2019). It can infest with great ease sound grains, but its development is benefited from the presence of frass or cracked particles (Kavallieratos et al., 2017a).

Habitat List

CategorySub categoryHabitatPresenceStatus
Terrestrial    

Biology and Ecology

Genetics

Trogoderma granarium genetics have been illustrated and used in separating this species from other species of the same genus, such as Trogoderma variabile or Trogoderma inclusum (Olson et al., 2014; Castañé et al., 2020). The mitochondrial genome of T. granarium, along with that of other Dermestidae has been recently published by Zeng et al. (2021).

Reproductive Biology

Based on what has been reported, the optimal temperatures for development of T. granarium range between 20 and 35ºC (Lindgren et al., 1955; Lindgren and Vincent, 1959), while at 35ºC, population growth of this species can be 10-250 times higher than Rhyzopertha dominica or Sitophilus oryzae (Kavallieratos et al., 2017a). However, with the exception of the adults that emerge from diapausing larvae, female adults of T. granarium lay less eggs than other major stored product insects (Hadaway, 1956; Karnavar, 1972; Odeyemi and Hassan, 1993). It is considered that T. granarium can have up to ten generations per year, if the conditions prevailing are suitable. Within the temperature range of 30 and 33ºC, the population increase of T. granarium has been calculated to be between 19 and 44 times per generation, corresponding to 2-2.5 times per week (EPPO, 2013; Athanassiou et al., 2016).
Adults of T. granarium do not feed and live less than 14 days (Gourgouta et al., 2021). Both sexes have developed wings, but they do not fly (Hadaway, 1956; Banks, 1977). Larval diapause is considered as the key characteristic that greatly contributes to its range expansion (Hadaway, 1956; Athanassiou et al., 2019). Diapause can last for several years, and can be interrupted for ‘foraging excursions’ (Burges, 1962a, b; Aitken, 1975; Bell, 1994; Wilches et al., 2016). T. granarium can have up to ten generations per year, and its population growth is extremely high at temperatures that are 30ºC or higher (EPPO, 2013; Kavallieratos et al., 2017a).
Physiology and Phenology

Longevity

Recent data has shown that egg hatch occurs within a few days after laying, usually 2-4 days (Gourgouta et al., 2021; Lampiri and Athanassiou, 2021). Egg to adult development time can be between 39 and 45 days at 30ºC, but can take 220 days at 21ºC (Athanassiou et al., 2019). Moreover, at temperatures lower than 30ºC, larval development can be interrupted, due to diapause induction (Wilches et al., 2016). Larval instars vary remarkably according to temperature and sex, but at temperatures of 30ºC or higher, development can be very rapid, 15 days, or even shorter (Hadaway, 1956; Burges, 1962a, b; Gourgouta et al., 2021; Lampiri and Athanassiou, 2021). Adults are short lived, and recent studies show that only a few can live more than 7 days, while all adults are dead before the completion of 14 days (Gourgouta et al., 2021).

Activity Patterns

As noted above, adults are not able to fly. This is important in the case of eradication planning as early detection may drastically reduce further spread. Also, adults can be continuously reproduced indoors, without access to the outside environment (Athanassiou et al., 2019). On the other hand, larvae are cryptic, can hide in cracks and crevices, and are usually detected when the species has reached high population densities.

Population Size and Density

At temperatures that exceed 30oC, T. granarium has a rapid population growth, and can easily outcompete other stored product insect species, especially in dry conditions (Kavallieratos et al., 2017a; Athanassiou et al., 2019). In laboratory studies with population growth in vials, the vast majority of the individuals were larvae (Athanassiou et al., 2016; Kavallieratos et al., 2017a; 2019).

Nutrition

Larvae can feed upon a wide variety of commodities and substances (Hagstrum and Subrananyam, 2009; Athanassiou et al., 2016; Kavallieratos et al., 2019). Recent studies show that young larvae of T. granarium are able to damage sound grain kernels (Athanassiou et al., 2016; Kavallieratos et al., 2017a).

Associations

Trogoderma granarium individuals have been found to be associated with numerous symbionts, such as Spiroplasma, and also Pseudomonas and Halospirulina, as well as Wolbachia (Li et al., 2015; Wilches et al., 2018).

Environmental Requirements

Dry conditions are key in the population growth of this species, where it can multiply remarkably fast (Barak, 1989; Kavallieratos et al., 2017a).
Larval development in T. granarium does not occur at temperatures below 21°C, but can proceed at very low humidity, for example at 25°C and 2% RH. Development is most rapid in hot, humid conditions, taking about 18 days at 35°C and 73% RH. Under these conditions, the average number of larval moults is four for males and five for females, although this is highly variable (Hadaway, 1956).
There are two genetic types of T. granarium larvae: those that are able to undergo a facultative diapause and those that are unable to do so. The former is stimulated to diapause by adverse conditions, such as extremes of temperature, humidity and crowding. When almost mature, the larvae leave their food and seek out a refuge such as a crevice in a store (Burges, 1962a, b; 1963). Their respiration then drops to an extremely low level. Large numbers of larvae in this condition may be found together and, although they are inactive, will seek a new refuge if disturbed. These larvae moult periodically with scarcely an increase in the rate of respiration. The larvae also feed occasionally; their respiration returns to normal during this time.
Without food, diapausing larvae may survive about 9 months; with food, they may live for 6 years. In this state of very low metabolic activity, they are extremely resistant to the effects of contact insecticides or fumigants; complete disinfestation may thus be difficult. The larvae leave diapause and pupate if subjected to a considerable temperature shock, i.e. a much lower temperature for at least a month, and then a return to warm conditions. A similar, but less effective, stimulus is the introduction of fresh food. Diapause seems to assist the larvae in surviving adverse conditions; it may also promote dispersal, as diapausing larvae are frequently found on movable objects or transport equipment such as sacks and lorries.
The pupa of T. granarium usually remains inside the skin of the final-instar larva. Pupal development is unaffected by humidity and varies in length from 5 days at 25°C to 3 days at 40°C. On adult emergence, the pupal skin is pushed to the posterior end of the larval skin; the adult remains within the skin for a day or more.
When the adults have fully emerged, copulation may take place immediately. To aid reproduction, the virgin females secrete a pheromone that attracts unmated males and, to a lesser extent, mated males and other females; reproduction sites may be established in this way. Female T. granarium only need to mate once. After copulation, oviposition commences immediately at 40°C and lasts 3-4 days, while at 25°C, there is a pre-oviposition period of 2-3 days, and oviposition may extend over 12 days. The abdomen of the newly emerged female is distended by its eggs and several segments may extend beyond the elytra; the abdomen returns to normal as oviposition takes place. Temperatures between 25 and 40°C seem to have little or no effect on the average number of eggs laid, which is approximately 35 per female. The females die soon after oviposition is complete; the males live 1-4 days longer. Under optimal conditions, T. granarium can sustain a rate of increase of 12.5 times per lunar month. The adults possess wings but have never been known to fly.

Air Temperature

ParameterLower limit (°C)Upper limit (°C)
Absolute minimum temperature1945

Seedborne Aspects

Effect on Seed Quality

The infestation of various types of seeds, such as grain kernels, by T. granarium is morphologically different to that of other major stored product insect species. Species like Rhyzopertha dominica and Sitophilus oryzae bore holes into the kernels, while T. granarium larvae can start the infestation from the external part of the kernel, turning the seed into dust. Loss of seed germination can occur in a very short period of time (Ahmedani et al., 2006).

Pathogen Transmission

As in the case of other stored product insect species, T. granarium can transfer several fungi to grain, causing deterioration and degradation (Kteo and Mohammed, 2019). Nevertheless, as this species can develop in conditions that are drier than those required for stored product insects, transfer of fungi may be more vigorous in other species, such as S. oryzae (Sinha and Sinha, 1990).

Notes on Natural Enemies

There are disproportionally few data for the utilization of predators and parasitoids for the control of T. granarium, as compared with other stored product insects. The warehouse pirate bug, Xylocoris flavipes, has been found to be associated with T. granarium larvae (Ahmed et al., 1991; Rahman et al., 2009). Predatory mites have been observed to use eggs of T. granarium as prey (Rahman, 1942). Several studies have documented the potential of using the parasitoids Laelius pedatus and Anisopteromalus calandrae (Kapil and Chaudhary, 1973; Ahmed, 1996; Al-Kirshi et al., 1997).
Regarding entomopathogens, the nematodes Steinernema carpocapsae and Steinernema masoodi have been successfully evaluated for the control of T. granarium larvae (Ali et al., 2011; Rumbos and Athanassiou, 2017). The entomopathogenic fungi Beauveria bassiana and Metarhizium anisopliae were pathogenic to khapra beetle adults and larvae (Bilal et al., 2017; Rumbos and Athanassiou, 2017). Bacillus thuringiensis was effective against larvae of T. granarium (Al-Hamdani et al., 2018).

Natural enemies

Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Acaropsellina doctaPredator
Eggs
    
Adelina triboliiPathogen
Adults/Eggs/Larvae/Nymphs/Pupae
    
Amphibolus venatorPredator
Adults/Eggs/Larvae/Nymphs/Pupae
    
Anisopteromalus calandraeParasite
Larvae/Pupae
    
Blattisocius keeganiPredator
Eggs
    
Dinarmus basalisParasite
Larvae/Pupae
    
Holepyris sppParasite
Larvae/Pupae
    
Pyemotes tritici (itch mite, straw (USA))Parasite
Larvae/Pupae
    
SynopeasParasite
Larvae/Pupae
    
SynopeasParasite
Larvae/Pupae
    
Theocolax elegansParasite
Larvae/Pupae
    

Impact

T. granarium is a serious pest of cereal grains and oilseeds, and many countries, including the USA, Australia, China, Kenya, Uganda and Tanzania, have specific quarantine regulations against possible importation. Massive populations of the insect may develop and grain stocks can be almost completely destroyed. Infestations of T. granarium are well known in large-scale stores but there appear to be no documented cases of infestations in farms.Losses due to T. granarium, sometimes in conjunction with other storage pests, have been reported in the literature and are summarized as follows.Losses in wheat grain stored in PVC bins after 90 days were 23.06% due to T. granarium, Tribolium castaneum, Sitophilus oryzae and Rhyzopertha dominica compared with 1.73% in fumigated bins (Singh et al., 1994).In a grain silo survey in Iraq between 1977 and 1978, T. granarium was present in more than 50% of samples. Infestation levels ranged up to 685 insects/kg grain. The mean percentage of infested grains ranged from 2.5 to 5.7% according to the origin of the wheat. The percentage wheat loss ranged from 3.1 to 6.6 (Al-Saffour and Kansouh, 1979).In Punjab, India, populations of T. granarium varied from 121 to 415 per 500g of wheat in a state survey in 1971-72. The pest damaged 9-14.5% of the grain resulting in 1.04-3.02% weight loss (Bains et al., 1976).In laboratory tests, feeding losses caused by infestations of T. granarium on wheat grain were estimated. The percentage infestation 30 days after the release of 10 pairs of adults into tubes containing 20g of grain was 9.4 and 15% at 30 and 36°C (optimum temp.), respectively. The percentage net loss in weight was 1.1 and 2.6, and percentage loss in viability was 12 and 24 at the different temperatures (Prasad et al., 1977).Analysis of wheat grain samples containing 5 to 100% T. granarium-infested grains showed that levels of protein, gluten, crude fat, ash, reducing and non-reducing sugars, and sedimentation value decreased with increased numbers of damaged grains. Values for alcoholic stability and free fatty acids increased. The proportion of seeds that germinated varied from nil when all kernels were damaged to 95% for no damage. Damage caused a loss in weight averaging 16.36% (Girish et al., 1975). T. granarium has also been shown to decrease the mineral content of maize (Jood et al., 1992).

Impact: Economic

Trogoderma granarium is a serious pest of cereal grains and oilseeds, and many countries, including the USA, Australia, China and Canada, have specific quarantine regulations against possible importation. Massive populations of the insect may develop and grain stocks can be almost completely destroyed (Athanassiou et al., 2016; Kavallieratos et al., 2017a). Losses due to T. granarium, sometimes in conjunction with other storage pests, have been reported in the literature and are summarized as follows.
Losses in wheat grain stored in PVC bins after 90 days were 23.06% due to T. granarium, Tribolium castaneum, Sitophilus oryzae and Rhyzopertha dominica compared with 1.73% in fumigated bins (Singh et al., 1994).
In a grain silo survey in Iraq between 1977 and 1978, T. granarium was present in more than 50% of samples. Infestation levels ranged up to 685 insects/kg grain. The mean percentage of infested grains ranged from 2.5 to 5.7% according to the origin of the wheat. The percentage wheat loss ranged from 3.1 to 6.6 (Al-Saffar and Kansouh, 1979).
In Punjab, India, populations of T. granarium varied from 121 to 415 per 500 g of wheat in a state survey in 1971-1972. The pest damaged 9-14.5% of the grain resulting in 1.04-3.02% weight loss (Bains et al., 1976).
In laboratory tests, feeding losses caused by infestations of T. granarium on wheat grain were estimated. The percentage infestation 30 days after the release of ten pairs of adults into tubes containing 20 g of grain were 9.4 and 15% at 30 and 36°C, respectively. The percentage net loss in weight was 1.1 and 2.6, and percentage loss in viability was 12 and 24 at the different temperatures (Prasad et al., 1977).
Analysis of wheat grain samples containing 5 to 100% T. granarium-infested grains showed that levels of protein, gluten, crude fat, ash, reducing and non-reducing sugars and sedimentation value decreased with increased numbers of damaged grains. Values for alcoholic stability and free fatty acids increased. The proportion of seeds that germinated varied from nil when all kernels were damaged to 95% for no damage. Damage caused a loss in weight averaging 16.36% (Girish et al., 1975). T. granarium has also been shown to decrease the mineral content of maize (Jood et al., 1992).
Athanassiou et al. (2016) reported that, at the laboratory scale, an initial number of 20 T. granarium individuals in 20 g of oats, triticale and wheat, resulted in a population of 128, 199 and 182 individuals after 60 days, respectively. Moreover, in that study, at the same interval, the population on whole barley flour, whole soft wheat flour and semolina, were 1458, 1020 and 996, respectively. Nevertheless, population growth of T. granarium in non-grain commodities was much more reduced in comparison with grains and related amylaceous products. In bags containing wheat stored under semi-field conditions, an initial infestation of the commodity with 100 larvae, increased more than ten times 8 weeks later, causing a significant increase in the numbers of insect damaged kernels (Scheff et al., 2021).

Impact: Social

Trogoderma granarium contaminates the substrates that it infests with body parts and cast skins, the consumption of which by humans can cause health disorders (Morison, 1925; Pasek, 1998; Stibick, 2007; Myers and Hagstrum, 2012). It is generally considered that the increase of the infestation frequency by T. granarium will threaten international food trade and global food security.

Risk and Impact Factors

Invasiveness

Proved invasive outside its native range
Has a broad native range
Capable of securing and ingesting a wide range of food
Has high reproductive potential

Impact outcomes

Damages animal/plant products
Negatively impacts trade/international relations

Likelihood of entry/control

Highly likely to be transported internationally accidentally
Difficult to identify/detect as a commodity contaminant
Difficult/costly to control

Detection and Inspection

Populations of T. granarium may be monitored using commercially available pheromone traps (Smith et al., 1992). However, the female-produced sex pheromone, which is a mixture of 92% of the Z and 8% of the E isomer of 14-methyl-8-hexadecenal (Cross et al., 1976), is also attractive for other species of this genus, such as Trogoderma glabrum, Trogoderma variabile and the larger cabinet beetle, Trogoderma inclusum. As adults do not fly, the most widely used trapping devices for the detection of T. granarium are floor or wall traps, that can also contain food attractants, such as bran. Moreover, the potential presence of different species of the genus Trogoderma in the same trapping unit might lead to misidentification (Athanassiou et al., 2019). Generally, the response of T. granarium adults to trapping devices is rather poor (Gourgouta et al., 2021). The female-produced sex pheromone of Trogoderma spp. has no adverse effects on the capture of Rhyzopertha dominica and Tribolium castaneum but captures of T. variabile are reduced when the T. castaneum pheromone is present (Dowdy and Mullen, 1998).

Prevention and Control

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.

Prevention

Due to its high phytosanitary importance, it is critical that introduction of T. granarium should be based on a proper phytosanitary control, especially in the commodities in which it is often detected (Myers and Hagstrum, 2012). In this regard, priority can be given in imports from khapra beetle-free geographical zones, based on phytosanitary and quarantine regulations. Proper monitoring at the entry ports is one of the key elements in prevention strategies towards the avoidance of establishment of T. granarium.

Eradication

There are successful eradication paradigms in the past, that resulted in constituting certain geographical zones as khapra beetle-free areas (Athanassiou et al., 2019). Once it is detected in a certain facility, the introduction record should be tracked, and the facility has to be closed, until it is proven that T. granarium has been totally eradicated. A zone around this facility, including adjacent facilities has to be regulated as well. Eradication programmes involve the application of fumigants and contact insecticides, and also a robust monitoring system.
Control

Chemical Control

Grain may be protected from T. granarium by the admixture of insecticides. T. granarium is, as far as it is known, susceptible to many of the contact insecticides normally used on stored grains, at concentrations that are effective for the control of other major stored product beetle species (Athanassiou et al., 2019). For instance, Kavallieratos et al. (2016; 2017b) found that different insecticides belonging to various groups with different modes of action were effective for the control of adults and larvae of T. granarium in both grains and on concrete surfaces. Grain stocks may be fumigated with phosphine to eliminate existing infestations, but these treatments provide no protection against re-infestation (Gourgouta et al., 2021; Lampiri and Athanassiou, 2021). If T. granarium is present, then fumigations should be undertaken for a longer than normal period, and might exceed 7 days, as diapausing larvae have low susceptibility to the fumigants. However, more recent published data clearly show that the egg of this species may be more tolerant to phosphine than the diapausing larvae. For instance, Gourgouta et al. (2021) shown that, after 3 days of exposure, adults, pupae, diapausing larvae and non-diapausing larvae could be totally controlled with 50 ppm of phosphine, while the respective concentration for eggs was 1000 ppm. In a more recent work, Lampiri and Athanassiou (2021) found that the age of the eggs of T. granarium was an important factor regarding tolerance to phosphine, while all eggs were dead after exposure at 750 ppm for 4 days. Nevertheless, there are confirmed cases of populations of T. granarium that are resistant to phosphine (Yadav et al., 2020).

Biological Control

Biological control has not been practised against T. granarium, under field conditions, despite the fact that there are data that show that certain biocontrol agents may be effective (Khashaveh et al., 2011; Athanassiou et al., 2019).

Cultural Control and Sanitary Methods

Good store hygiene plays an important role in limiting infestation by T. granarium. The removal of infested residues from the previous season's harvest is essential, as is general hygiene in stores; all spillage should be removed and all cracks and crevices filled.

Extreme temperatures and other non-chemical control methods

Recently published works have shown that acclimatized diapausing larvae of T. granarium are tolerant to both heat and cold. Wilches et al. (2019) indicated that in order to totally control all life stages of T. granarium (incl. diapausing larvae), a 2-hour exposure is needed at 60ºC. Similarly, diapausing and acclimatized larvae required exposure to -10ºC for 87 days for complete mortality (Wilches et al., 2017).
Low oxygen, as well as high CO2 can control T. granarium, and supress its progeny production capacity (Vassilakos et al., 2019).
Both larvae and adults of this species have been found susceptible to diatomaceous earths, at concentrations that are also effective for other major stored product insect species (Kavallieratos et al., 2017b).

Links to Websites

NameURLComment
GISD/IASPMR: Invasive Alien Species Pathway Management Resource and DAISIE European Invasive Alien Species Gatewayhttps://doi.org/10.5061/dryad.m93f6Data source for updated system data added to species habitat list.
Global register of Introduced and Invasive species (GRIIS)http://griis.org/Data source for updated system data added to species habitat list.

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