Regulatory Control of Khapra Beetle

The following list of references includes many from Myers and Hagstrum 2013 and additional references, websites and annotations are included to expand upon their summary. Several studies predict possible establishment of the khapra beetle, Trogoderma granarium, based upon climate (Banks 1977 for Australia, Sinha 1963 for Canada, Bahr 1966 for East Germany, Dal Monte 1960 for Italy, Mitsui 1965 and Sinha and Utida 1967 for Japan, Howe and Lindgren 1957 and Howe 1958 for U.S.A.). The suitability of climate in Iran (Freeman 1958a), Nigeria (Howe 1952), Pakistan (Freeman 1958b) and Turkey (Freeman 1962) for Trogoderma granarium has also been considered. Some of the references are to research supporting regulatory effort or to educational materials prepared to help involve public in regulatory programs. Additional relevant references can be found on Life History and Population Ecology pages.

Khapra beetle destroys more commodity than it consumes. Khapra in Hindi and Urdu means “the destroyer”. Khapra beetles have wings but cannot fly. Larval diapause allows it to survive for several years without food. Diapausing larvae tend to remain hidden, but may come out of hiding to feed. It feeds on many commodities and is more tolerant than other species to many pesticides and extreme temperature and relative humidity, preferring high temperatures and low relative humidities for larval development. In Australia, Trogoderma variabile was not eradicated as easily as Trogoderma granarium because it was able to disperse by flight (Butcher 1994, Castalanelli et al. 2011, Emery 1999) and a strategy of containment was adopted.

Several dichotomous keys using adult and larval morphology for species identification are available (Anonymous 1955, 2016, Banks 1994, Barak 1995, Beal 1956, 1960; Kingsolver 1963, Mawlood and Abdul-Rassoul 2000, Okumura 1955, 1966, Okumura and Blanc 1955, Zhongping et al. 2017, Zhang et al. 2008) and there have been several studies developing molecular identification methods (Chen et al. 2010, Khidr et al. 2017, Ma et al. 2009, Olson et al. 2014, Shen 1994, Stuart et al. 1994, Yulin et al. 1999, Zhang et al. 2017, Zheng et al. 2016). There are generally more larvae than adults to be identified because larvae diapause and may be in the larval stage for several years while adults are short lived. Burges (1959b) found that insects in larval stage for more than 7 weeks are in diapause. Trogoderma granarium is a species with data in Barcode of Life Database (BOLD) software in the GenBank but this data has not been open to the public (Boykin et al. 2012). Further field validation of the reliability of the molecular identification methods identifying species are likely to be needed before these methods are used.

Six lists of locations (1977 to 2016) at which khapra beetle is reported as currently or previously present or intercepted in imported commodities are compared in Locations with Khapra table. Myers2012 list includes only records for imports into the US and 43 locations. Australia2016 includes 75 locations. The second column of table shows the number of lists in which a location is included. Twenty six percent of 129 locations are on 4 or 5 of lists and 40 % are on only 1 or 2 lists. Yugoslavia may include Croatia, Serbia and Slovakia, and United Kingdom may include England and Ireland. EPPO2016 and Banks1977 list include 75 and 59 locations and categorize all locations while French2005 list includes 90 locations, but does not categorize all (location name is given when not categorized). SPIR2009 list includes 89 locations. The Central or South American countries, Argentina, Bolivia, El Salvador and Uruguay are only on 1 or 2 lists, but Venezuela is on 4 lists. Banks1977 and French2005 provide literature citations for the sources of records.

Before containerized shipping was introduced in the late 1960’s, cross infestation between cargos in a ship’s hold and infestation of commodities from residual insect infestations in ship’s hold were a problem (Hurlock 1962). Shipping containers should reduce the cross infestation problem, but carryover of residual infestations in shipping containers is now a problem. Many of khapra beetle interceptions are probably a result of shipping containers used for agricultural products later being used for non-agricultural products such as screws, rain ponchos, wood veener and automobiles. When khapra beetle infested residues remain undetected in empty shipping containers, a khapra beetle infested shipment may originate anywhere in the world that these shipping containers are used.

The following table sorts locations by their relative importance as sources of khapra beetle infestation in international trade based upon comments from CABI and EPPO. The table also provides a chronology of some of the expansion of khapra beetle geographic distribution. Locations with Khapra table above has an additional 50 locations not in this table. No. Lists column refers to the number of lists in Locations with Khapra table above that included a location.

Sources of khapra beetle, Trogoderma granarium, infestation1,2


Location                           Relative importance                               No. Lists


Algeria                                 Widespread, introduced                                        5

Cyprus                                 Widespread                                                               6

Morocco                              Widespread, introduced                                         5

Syria                                     Widespread                                                               6

Tunisia                                 Widespread, introduced                                        6

Zimbabwe                             Widespread, found 1955*                                       5

Afghanistan                       Present                                                                      5

Bangladesh                        Present                                                                      5

Burkina Faso                     Present, introduced                                               5

Egypt                                     Present, introduced, found 1935*                       6

India                                     Present, native                                                         6

Iran                                       Present                                                                      6

Iraq                                       Present                                                                      6

Korea, Republic                   Present                                                                      5

Lebanon                                Present                                                                      5

Libya                                    Present                                                                      5

Mali                                      Present, introduced                                                6

Mauritania                        Present, introduced                                                5

Myanmar                           Present                                                                      5

Niger                                    Present, introduced before 1943                          5

Pakistan                              Present                                                                      6

Saudi Arabia                     Present                                                                      5

Senegal                                Present, introduced before 1943                          6

Somalia                                  Present                                                                      4

Sri Lanka                            Present                                                                      6

Sudan                                   Present, introduced                                                6

USSR, former                       Present, introduced                                                 .

Zambia                                  Present, introduced 1954*                                      6

Zanzibar                                Present, introduced 1956*                                      2

Israel                                    Restricted distribution, introduced 1939             6

Nigeria                                 Restricted distribution, intro. before 1943          6

Spain                                      Restricted distribution, introduced                       5

Switzerland                           Restricted distribution                                             3

Turkey                                  Restricted distribution, introduced 1954             6

Yemen                                    Restricted distribution                                              5

Angola                                   Absent, unreliable record                                          5

China                                     Absent, unreliable record                                          5

Cote d’Ivoire                         Absent, unreliable record                                          5

Gambia                                  Absent, unreliable record                                          5

Guinea                                   Absent, unreliable record                                           5

Italy (Sicily)                           Absent, unreliable record                                          1

Malaysia (peninsular)          Absent, unreliable record                                         1

Mozambique                          Absent, unreliable record                                         5

Philippines                             Absent, unreliable record                                         5

Reunion                                  Absent, unreliable record                                         4

Serbia-Montenegro              Absent, unreliable record                                          3

Thailand                                 Absent, unreliable record                                          5

Belgium                                 Absent, formerly present                                            3

Denmark                               Absent, formerly present                                            4

Indonesia                               Absent, formerly present                                           4

Ireland                                   Absent, formerly present, found 1948*                    4

Kenya                                     Absent, formerly present                                            4

Luxembourg                         Absent, formerly present                                            3

Netherlands                          Absent, formerly present                                            3

New Zealand                        Absent, formerly present                                             3

Russian Federation             Absent, formerly present                                            4

Sierra Leone                          Absent, formerly present                                            5

Sweden                                   Absent, formerly present                                            3

Taiwan                                    Absent, formerly present                                            5

United Kingdom                   Absent, formerly present, prior 1908*                    2

Uruguay                                 Absent, confirmed by survey                                      2

Venezuela                              Absent, formerly present                                            4

Australia                                Eradicated, introduced 2007, 2016*                        5

Germany                                Eradicated, found 1921                                               4

Hungary                                 Eradicated                                                                     3

Japan                                      Eradicated, found 1923, absent 1950, intro 1964*  5

Malaysia                                 Eradicated                                                                     5

Mexico                                    Eradicated*                                                                   4

South Africa                          Eradicated, found 1954*                                             4

Tanzania                                Eradicated, introduced 1960*                                    6

USA                                         Eradicated, introduced 1946*                                    4

Austria                                    Absent, intercepted only, confirmed by survey      3

Bulgaria                                  Absent, intercepted only                                             3

Croatia                                   Absent, intercepted only                                              3

Czech Republic                    Absent, intercepted only                                              3

Italy                                       Absent, intercepted only, found 1953*                      4

Poland                                   Absent, intercepted only                                              3

Portugal                                 Absent, intercepted only                                             4

Slovakia                                Absent, intercepted only                                              3


1. Modified from CABI Invasive Species Compendium at: (Last modified 11 October 2017). Comments and countries included are essentially the same as those in EPPO Global Data Base. Trogoderma granarium at:

2.  Twenty-five countries recognized by US as harboring endemic populations of khapra beetle (in   bold) are from: Stibick, J., 2007. New Pest Response Guidelines: khapra Beetle. USDA–APHIS–PPQ–Emergency and Domestic Programs, Riverdale, Maryland. Available from: URL: http:// * Indicates from another source.

The host range for the khapra beetle, Trogoderma granarium, has been studied in two ways: by recording the commodities infested and by determining in the laboratory whether commodities are suitable for their development. Khapra infested_suitable commodities table shows that 31 commodities have been studied using both methods. Multiple infestation records (2-6) are available for many commodities and more records are available for grains (maize 7, rice 12, wheat 12) and peanuts (11). The majority of these commodities (20) were suitable for development in 1-3 studies or for wheat in 6 studies. In a few cases (4) commodities were suitable in one study but not another. The remaining 7 commodities although found to be infested were not suitable for development in laboratory studies. Another 110 commodities have been studied by one of the two methods. Twenty six of these commodities were found to be suitable for development in the laboratory. The final 44 commodities in the list have been shown to be unsuitable for development in the laboratory.

The Atlas and SPIR together had 104 host commodities and the other five publications, each with 27-38 hosts, added 38 new hosts that were not in the Atlas or SPIR (Host of khapra table). Overall, the 143 hosts were in 22 of the 28 commodity categories in the Atlas (p. 201). Fifteen of the hosts were in 4 to 6 of the publications and these hosts included some of the most prevalent commodities in the marketing system. The majority of these were grains (6) or pulses (6). A little more than half (74) of the host commodities were only in one of the six publications. About one third (27) of these were seed and 22 were from 1955 to 2005 publications. CERIS2004 called several of hosts grocery items (groc) and indicated that khapra beetle could not fully develop on 4 hosts. Strong1959 classified host commodities as primary (prim), secondary (sec) or miscellaneous (misc).

The following table has information from 62 press releases for khapra beetle interceptions between 2010 and 2018 (United States Customs and Border Protection web site, ) and 12 additional interceptions in 2012 (Mason 2013). The number of interceptions included was highest with 16 in 2011 and 19 in 2012, and ranged from 2-11 for the other years. More interceptions were on rice (32) and bean (9) than any other commodity. Khapra beetle was intercepted 1-3 times on each of an additional 21 commodities, many of them legumes. Rice was the only cereal grain. Nearly half of the interceptions were in airline passenger luggage. Interceptions on screws, rain ponchos and wood veener were probably a result of the carryover of khapra beetle infestations in shipping container from previous cargo. The most interceptions were for commodities from India (19) Saudi Arabia (18), Sudan (11) and Pakistan (7). There were 1-6 interceptions from 9 additional countries.

Interceptions of Trogoderma granarium by year1. ___________________________________________________________

Month            Point Interception                Commodity Infested              Country of Origin

___________________________________________________________        2010

March             New York*                              dried beans                              India

May                 Oakland, CA                           organic brown rice

July                  Oakland, CA                          rice

August            Houston, TX*                        rice                                            Saudi Arabia


January           Los Angeles, CA                   rice                                           Saudi Arabia

January           Los Angeles/LB, CA           basmati rice                            Pakistan

January           St. Croix, Puerto Rico        cargo vessel

June                Port Huron, MI*                 decorative fan                          India

June                 NY/NJ                                  rice                                            Pakistan

July                  Indianopolis, IN                 barley seed                              India

August            Chicago, IL                            rice                                           India

August             NY/NJ                                   rice                                           Pakistan

September      NY/NJ                                   safflower seed                        India

September      Los Angeles, CA                  rice

October           Houston, TX                       melon seed                             Saudi Arabia

October           Norfolk, VA                         safflower seed                        India

October          Los Angeles, CA                  basmati rice                            India

October          Los Angeles/LB, CA           beans                                       UAE

November      Los Angeles, CA                  rice                                           India

December      New York, NY                     dried seeds, roots, spices     Saudi Arabia


January         San Francisco, CA*            pumpkin seed                         Saudi Arabia

January         San Francisco, CA*            cumin seed                              Saudi Arabia

February       New York, NY                      food                                          UAE

February       Detroit, MI*                         pepper seed                            Iraq

February       Chicago, IL*                         rice                                          Saudi Arabia

February       Chicago, IL                           rice                                          Saudi Arabia

February       Cincinnati, OH                    rice                                           Saudi Arabia

February       Dallas, TX                            rice                                           Saudi Arabia

February       Dallas, TX*                          bean                                         Sudan

March            New York, NY                    moong dal                               UAE

March            New York, NY                    Urad dal(split black lentil)  UAE

March            Phoenix, AZ*                      rice                                           India

April               Detroit, MI Cargo              wood veener                          Ivory Coast

July                Charleston, SC                    soybean                                  India

August           Chicago, IL                         rice

September    Pembina, ND                      outside bag foodstuff

October          Philadelphia, PA*              rice                                         Saudi Arabia

October          San Diego, CA*                  lentils                                     India

December      Norfolk, VA                        rice                                          UAE


February        Baltimore, MD                  celery seed

May                 Dallas, TX*                        seed, dried bean                   Sudan, India

August            Chicago, IL                         rice                                         Pakistan

September     Pittsburgh, PA*                 rice                                         Saudi Arabia


August            Abu Dhabi, UAE*              chickpea                                Pakistan

August            Baltimore, MD                    rice                                        Pakistan

August            Washington, DC*               rice                                        Saudi Arabia

September      Atlanta, GA*                      pigeon peas                          Sudan

September      Philadelphia, PA*             rice                                        Saudi Arabia

September      Philadelphia, PA*             acacia seed                          Sudan

September      Philadelphia, PA*              fava beans                          Sudan

November      Lewiston Bridge, NY         rain ponchos                      China


February         Dallas, TX*                         rice                                        India

February         Dallas, TX*                         fava bean                             Sudan

April                Dallas, TX*                          bean                                     Sudan

June                San Francisco, CA             chickpea                               India

June               Chicago, IL*                        dried hibiscus leaves         Iraq

July                Phoenix, AZ*                       rice                                        Saudi Arabia

July                 Chicago, IL*                       peanuts, rice                       India

August            Toronto, Canada*             bean                                     Somalia

November      Norfolk, Va                        organic soybeans               India

November      Norfolk, Va                        split lentils and spices      UAE

December      Dublin, Ireland*               chickpea peanut mix         India


March              Baltimore, MD                  cumin seed                        India

May                  Dallas, TX*                        seeds and dried beans     Sudan, India

September      Philadelphia, PA               rice                                      Saudi Arabia

December       Port Huron, MI                 moong dal                          India, Nepal

December       Atlanta, GA*                      bean                                     Nepal


January           New York, NY                  rice                                        Saudi Arabia

May                  Baltimore, MD                 screws                                  Thailand

May                  Houston, TX*                  hibiscus and chickpeas     Sudan

June                 Newark, NJ                      rice                                        Bangladesh

August             Los Angeles, CA               rice                                       Pakistan

August             Philadelphia, PA*            dried berries                       Sudan

October           Dallas, TX*                        jujubes                                 Sudan


January          Washington, DC*              basmati rice                        Saudi Arabia

March             Washington, DC*               cowpea                                Nigeria


1 Interceptions in luggage have * after point of interception. Trogoderma granarium were intercepted during preclearance in Abu Dhabi, Dublin and Toronto.


Pest Risk Assessment for the State of Oregon Trogoderma granarium (Everts) – Khapra Beetle

Poster: U.S. Agricultural Alert for international travelers departing from India

Agarwal, Manjree, Thamer Al-Shuwaili, Anupiya Nugaliyadde, Penghao Wang, Kok Wai Wong, and Yonglin Ren. 2020. Identification and diagnosis of whole body and fragments of Trogoderma granarium and Trogoderma variabile using visible near infrared hyperspectral imaging technique coupled with deep learning. Computers and Electronics in Agriculture 173, 105438.

Ahmedani, M. S., A. Khaliq, M. Tariq, M. Anwar, and S. Naz. 2007. Khapra beetle (Trogoderma granarium Everts): a serious threat to food security and safety. Pak. J. Agri. Sci. 44: 481-490.

Ahmdani, M. S. 2009. Phytosanitary Management of Trogoderma granarium Everts with Methyl Bromide Alternatives To Ensure Food Security And Safety. PhD diss., Department of Entomology, Faculty of Crop and Food Sciences, Pir Mehr Ali Shah (PMAS)-Arid Agriculture University, Rawalpindi, Pakistan.

Alkan, B. 1962 Die wichstigsten vorratsschadlinge in den Turkei. (The main stock pests in Turkey) Verh. XI Int. Kongr. Ent. Wien 1960. 2, 316-320. (Trogoderma granarium Evert found wheat, barley and other cereals, as well as flax and cottonseed). This pest has only been known in Turkey since 1954 and has hitherto been found in the provinces of Diyaribakir, Hatay, Urfa and Mardin; khapra does a lot of damage.)

Anonymous. 1955. Have you seen this in your grain, how to look for khapra beetle. USDA PA-261

Anonymous. 1955. Khapra beetle. A Situation Report USDA, ARS 22-17 (Aug) (Hearing on the advisability of a Federal quarantine on the khapra beetle was held at Denver, Colorado, December 1, 1954 resulting in a February 21, 1955 Federal quarantine order for the states of Arizona, California and New Mexico. A preliminary survey of 492 warehouses, milling and grain-handling establishments in 10 states – AZ, CA, CO, ID, NV, NM, OR, TX, UT, WA – from early April until mid-June found khapra beetles in 25 warehouses (Padget 1954). The beetle had been shipped from Arizona to New Mexico in sorghum seed. Additional survey of 4,500 properties in AZ, CA, CO, LA, NM, OK, TX and other states, found khapra beetles in 63 in AZ, 158 in CA (of which at least 45 made interstate shipments) and 4 in NM.

Shipping records of the past two years were examined for shipments likely to carry infestation. Inspections were made at destinations and railway cars carrying bulk grain. Four hundred cars were inspected. One dealer whose warehouse was infested with khapra beetle shipped to 30 states in the US and 3 in Mexico.)

Anonymous. 1955. Illustrated key to species of trogoderma and to related genera of dermestidae commonly encountered in stored grain in California. Calif. Dept. of Agriculture, Sacramento

Anonymous. 1960. A summary of information about the khapra beetle. USDA, Agricultural Marking Service, Marketing Quality Research Division, Stored-Product Insect Branch 390.

Anonymous 1960. Khapra beetle threatens nation’s stored grain and seed. USDA Picture Story 126

Anonymous. 1961. The khapra beetle, a pest of stored grain and cereal products. USDA PA-436. (Khapra beetle was found in >200 establishments in Arizona, >300 in California, 19 in New Mexico, 22 in Texas and 84 in Mexico. For eradication, buildings were wrapped with gas tight tarpaulins and fumigated with 5 lbs of methyl bromide per 1,000 cu ft for 48 hours. The area surrounding the structures was treated three times at 3-7 day intervals with a spray containing 5 lbs of malathion in each 100 gallons of diesel fuel. The sprayed area was raked or harrowed between applications to assure wetting of all debris and other materials.

When infested premises were treated in this manner, they can be released from regulation immediately. If infestations are light, other approved procedure may be used. Only part of the structures on infested premise may be treated or structures may be treated without draping with tarpaulins. If the modified procedure is used, the infested premise must remain under regulation for a year. Three inspections at least 90 days apart with the final during the last month of period must show that the premises are free of infestation before they can be released.)

Anonymous 1978. Khapra beetle (Trogoderma granarium). USDA.  Cooperative Plant Pest Report 3(10): 76. The first active infestation by Trogoderma granarium (Everts) in the eastern USA was detected in a warehouse at Linden in New Jersey in February 1978, where 2 larvae and some cast skins were collected from wood-borer holes in imported wooden crates containing wooden screens; additional larvae, cast skins and 1 dead adult were subsequently found at large in the same warehouse. Fumigation of the warehouse and delimiting surveys are planned. The last active infestion by T. granarium in the USA occurred in the west of the country in 1966.

Anonymous. 1981. United States – khapra beetle infestations reported. FAO Plant Protection Bulletin 29: 30.

Anonymous. 1983. Pests not known to occur in the United States or of limited distribution, No. 30: Khapra Beetle, USDA, APHIS, PPQ, Hyattsville, MD. pp. 1-11.

Anonymous. 2011. Plant protection profiles from Asia-Pacific countries (2009-2010). Asia and Pacific Plant Protection Commission (APPPC) and Food and Agricultural Organization (FAO). (Khapra beetle was intercepted 248 times, particularly on imports of coconut oil-cake from Indonesia, and in wheat bran from Sri Lanka entering Vietnam during 2009. In 2010, khapra beetle was intercepted 189 times on corn, soya bean, barley and millet imported from India. Khapra beetle was intercepted in an additional 104 shipments from India by early 2011.)

Anonymous 2016. ISPM 27 Diagnostic protocols for regulated pests DP 3: Trogoderma granarium Everts. International Plant Protection Convention (IPPC). (english); (spanish)

APHIS 1972. The khapra beetle : a pest of stored grains and cereal products. USDA Program aid 436.

Arizona Cooperative Report. 1954. A dermestid (Trogoderma granarium) – Arizona. USDA Coop. Econ. Ins. Rpt. 4(7): 131. (Feb 19) (Khapra beetle found at four seed storage or flour mills in Phoenix, two in Glendale, one Casa Grande and one in Tucson largely from buildings in which flour and trash accumulate.)

Armitage, H. M. 1954. Insect eradication procedures in incipient infestations. J. Econ. Entomol. 47(1): 6-12.

Armitage, H. M. 1955. Fumigation-eradication test on khapra beetle. Down to Earth 10(4): 2-3.

Armitage, H. M. 1955. Khapra beetle eradication fumigation test No. 1 – Imperial, Calif., Empty grain storage warehouse, January 1955. State of California Department of Agriculture Circular E-59 of April 15, 1955

Armitage, H. M. 1955. Khapra beetle fumigation test No. 2 – Alpaugh, Kings County, Calif., Grain storage warehouse containing bulk grain – Using forced gas recirculation. State of California Department of Agriculture Circular E-59-1 of June 17, 1955

Armitage, H. M. 1955. Khapra beetle fumigation test No. 3 – Corooran, Kings County, Calif., Grain storage warehouse containing bulk grain – Without use of forced gas recirculation.  State of California Department of Agriculture Circular E-59-2 of Aug 30

Armitage, H. M. 1958. The khapra beetle suppression program in the United States and Mexico. Pages 89 98 in: Proc. Int. Congr. Entomol., 10th. (Montreal, Canada) E. C. Becker, Ed. Mortimer Limited, Ottawa, Canada. (Trogoderma granarium was introduced in California before 1946 but incorrectly identi­fied as Attagenus piceus (now black carpet beetle, Attagenus unicolor) and spread to 16 counties in California, five in Arizona, and three in New Mexico before being correctly identified in 1953. By 1958, 51,000 premises in 27 states had been inspected, and the pest was eradicated in the United States by 1966.)

Arrow, G. J. 1917. The Khapra beetle (Trogoderma khapra, sp. n.) and Indian grain pest. Ann. Mag. Nat. Hist. 19: 481-482. RAE A 5: 359. (Khapra beetle was found in many cargoes of wheat from Karachi and Bombay into England, but had not occurred in grain imported from countries other than India, nor had it been known to establish in Europe.)

Bahr, J. 1966. Unter welchen Umständen sind in der Deutschen Demokratischen Republik Schäden durch den Khaprakäfer (Trogoderma granarium Ev.) zu erwarten? (Under what circumstances in the German Democratic Republic is damage by the khapra beetle (Trogoderma granarium Ev.) to be expected?) . Wiss. Z. Univ. Rostock, Mathem. -naturwiss. Reihe 15: 327-335.

Bailey, S. W. 1958. Position of the khapra beetle in Australia. FAO Plant Prot. Bull. 6 (5): 1-2. (A survey over a wide area of Australia found 30 species of dermestid but not khapra beetle. Except for the single record from Tasmania on a cargo of barley from South Australia, khapra beetle had never been found within Australia. All suspected infestations have been associated with shipping. In 1956, khapra beetle was intercepted in the UK in a shipment of barley from South Australia on a ship that 2 years earlier had carried a cargo infested with khapra beetle.)

Baker, J. L. and T. J. Ward. 1935. Note on the occurrence of the khapra beetle in malt lofts. Journal of the Institute of Brewing 41: 375. (When the steam pipes beneath the lofts were insulated khapra beetle was no longer a problem, although sacks infested with khapra beetle larvae were occasionally found.)

Banks, H.J. 1977. Distribution and establishment of Trogoderma granarium Everts (Coleoptera: Dermestidae): climatic and other influences. Journal of Stored Products Research, 13: 183–202. (Trogoderma granarium is absent from Australia though some parts are apparently climatically suitable. It is concluded that climatic factors at the ports may have prevented the establishment and subsequent transport of sufficient numbers to more climatically favourable areas inland. Prevailing temperatures at the ports appear adequate for T. granarium but the high relative humidity there seems to be the main controlling factor.)

Banks, H. J. 1994. Illustrated identification keys for Trogoderma granarium, T. glabrum, T. inclusum and T. variabile (Coleoptera Dermestidae) and other associated with stored products. CSIRO Australian Division of Entomology Technical Paper No. 32. 66 pp.

Barak, A. V. 1989. Development of a new trap to detect and monitor khapra beetle (Coleoptera: Dermestidae). Journal of Economic Entomology 82: 14701477.

Barak, A. 1995. Identification of Common Dermestids, pp. 187-196. In V. Krischik, G. Cuperus and D. Galliart (eds.), Stored Product Management, vol. E912. Oklahoma State University, Stillwater, Oklahoma.

Beal, R. S. 1956. Synopsis of the economic species of Trogoderma occurring in the United States with description of a new species (Coleoptera: Dermestidae). Annals of the Entomological Society of America. 49: 559-566.

Beal, R. S. 1960. Descriptions, biology, and notes on the identification of some Trogoderma larvae (Coleoptera, Dermestidae). U.S. Department of Agriculture, Agriculture Research Service Technical Bulletin No. 1228.

Belskaya, N. M. and L. G. Popova. 1978. Injurious insects in cargoes from India. Zashchita Rastenii 2: 42-43. (Trogoderma granarium was intercepted from Indian grain cargoes 188 times at port of Odessa, Ukraine between 1967 and 1976 (mainly 1973-1975) constituting 45% of all insect pest interceptions.)

Biosecurity New Zealand. 2007. Import risk analysis: Vehicle & machinery. Ministry of Agriculture and Forestry, Wellington, New Zealand.

Botha, John, Andraz Szito, Robert Emery, Mike Grimm, and Darryl C. Hardie. 2005. Grains Industry Biosecurity Plan Threat-Specific Contingency Plan. Plant Health Australia, Canberra ACT.

Boykin, Laura M., Karen Armstrong, Laura Kubatko, and Paul De Barro. 2012. DNA barcoding invasive insects: database roadblocks. Invertebrate Systematics 26(6): 506-514.

Burges H. D. 1959a. Dormancy of the khapra beetle: quiescence or diapause. Nature 184: 17411742.

Burges H. D. 1959b. Studies on the dermestid beetle, Trogoderma granarium Everts II. The occurrence of diapause larvae at a constant temperature, and their behaviour. Bulletin of Entomological Research 50: 407–422. (Mostly fourth instar or later instar larvae and a few third instar prefer 1.2 mm crevice to wider or narrower. The larvae entering crevices that pupated varied from 22% near hot air chamber to 69% in bin, below bubble on kiln wall. The remaining larvae resting in diapause moult occasionally and at long and irregular intervals emerge, presumably to replenish their food reserves by feeding and then pupate or hide again.)

Burges H. D. 1959c. Studies on the dermestid beetle, Trogoderma granarium Everts III. Ecology in malt stores. Annals of Applied Biology 47: 445–462. (Khapra beetle has been introduced into many times into British maltings since about 1908, imported in barley from the hot dry regions extending from India to the shores of the Mediterranean. It has been spread within Britain in malted barley, sacks and transport vehicles and has breed rapidly in malt stores. Control by cooling malt before storage in promising.)

Burges H. D. 1960. Studies on the dermestid beetle, Trogoderma granarium Everts IV. Feeding, growth, and respiration with particular reference to diapause larvae. Journal of Insect Physiology 5: 31734.

Burges H. D. 1962. Studies on the dermestid beetle, Trogoderma granarium Everts V. Reactions of diapause larvae to temperature. Bulletin of Entomological Research 53: 193213.

Burges H. D. 1963. Studies on the dermestid beetle, Trogoderma granarium Everts VI. Factors inducing diapause. Bulletin of Entomological Research 54: 57187.

Butcher, M. J. 1994. Decision making in regulatory entomology: the case of Trogoderma variabile in Western Australia. p. 1169-1172. In E. Highley, E. J. Wright, H. J. Banks and B. R. Champ (eds.), Stored Product Protection, Proceedings of the 6th International Working Conference on Stored-Product Protection, 17-23 April 1994, Canberra, Australia. CAB International, Wallingford, United Kingdom

Castalanelli, M. A., K. M. Mikac, A. M. Baker, K. Munyard, M. Grimm, and D. M. Groth. 2011. Multiple incursions and putative species revealed using a mitochondrial and nuclear phylogenetic approach to the Trogoderma variabile (Coleoptera: Dermestidae) trapping program in Australia. Bull. Entomol. Res. 101: 333-343.

Chen Y., Zhu S.F., and Chen K., 2010, A molecular identification of Trogoderma granarium based on partial COI gene, Zhiwu Jianyi (Plant Quarantine), 24(1): 22-23

Cotton, R. T. 1954. Khapra beetle…newcomer pest. Agricultural Situation 38(12): 3.

Cotton, R. T. 1955. Khapra beetle. Modern Sanitation 7(5): 26-28, 47-48 (May).

Cotton, R. T. 1955. What are we doing about the khapra beetle. Presented at the 10th Annual Conference, North Central Branch, Entomological Society of America, East Lansing, Michigan, March 24-25, 1955. (At the invitation of the Directors of the Western State Experiment Stations, the Stored-Product Insect Section of the Agricultural Marketing Service established a field station at Mesa, Arizona in August 1954 to conduct research on the khapra beetle under the direction of G. H. Spitler.

The Western States Cooperative Regional Project WM-16 was already set up for Maintaining Grain Marketability by Insect Control in Storage so this project was broadened to include a study of the khapra beetle, and a subcommittee headed by A. E. Michelbacher of California was established to coordinate research on this insect by the USDA, Arizona and California.)

Cotton, R. T. 1960. Let’s kill the Khapra beetle invader. Northwestern Miller, 264(8), 42-44. (Perez Simmons in February 1939 received a khapra beetle from wheat in Madera County California. Larvae seek hiding places for aestivation in many nonfood commodities so they can be found in an endless variety of products.  If allowed to spread, khapra beetle could be a major pest of seed establishments, feed mills, malting houses, flour mills, powdered milk factories, cereal plants, grain elevators, nut storage and packing plants, warehouses handling dried food products and in farm grain storage.

During 1954 and January 1955, quarantine measures were adopted by the states of AZ, CA, CO, GA, KS, LA, NV, NM, OK, OR, TX, UT and WY. By October 1960, the number of properties found and fumigated since the beginning of the program in 1955 are: Arizona 242, California 341, New Mexico 19, Texas 28 and Republic of Mexico 87 representing 170, 945, 254 cu ft. All of the formerly infested properties except three in Texas have been released from quarantine.

During the period of July 1, 1959 to June 30, 1960 the pest was intercepted 133 times on 82 ships with approximately 58, 900 inspections by Plant Quarantine Division. Subsequently, khapra beetle was found in imported peanuts in New York and through June 24, 1960, 55 cargoes have been found to be infested involving 45 different ships.

Khapra beetle ntercepted at ports of entry on gum karaya, gum Arabic, seed lac, baled bagging, crude guar meals and hulls, cotton card strips, tumeric roots, baled crepe rubber, automobiles, steel wire, steel beams, kegs of bolts, metal drums, soiled linen, oil paintings, rice, myrobalan, nuts, peanuts, cumin, poppy seeds, guar embryos and dried animal skins.)

Carncross, L. B. 1961. How we controlled Khapra beetle in burlap-wraped steel from Europe. Pest Control 29(10): 46, 50-52.

Curl, L. F. 1960. Domestic quarantine notices (Plant Pest Control No. 612, 27th Revision). Title 7 Agriculture Chapter III Part 301 Subpart khapra beetle, U.S. Department of Agriculture (Lists warehouses, mills and other premises in which khapra beetle detected and designated as regulated areas.)

Curl, L. F. 1964. Mexico-United States cooperative plant pest programs. J. Econ. Entomol. 57: 450-452.(Khapra beetle infestations reported in Baja Califor­nia in December 1954 were apparently from infested products originating in the United States. In Mexico total of 25 million cubic feet of storage space at 92 properties infested by khapra beetle were successfully fumigated with methyl bromide by September 1961.)

Dal Monte, G. 1954. Un nuovo nemico nei nostrimagazzini. (A new enemy in our warehouses) Molini d’Italia 5: 198-201.

Dal Monte, G. 1960. Il pericolo della diffusione in Italia del Trogoderma granarium, Everts. (The danger of the spread in Italy of Trogoderma granarium, Everts), pp. 313-316. In Anonymous Proceedings of the 11th International congress of entomology, 17-25 Aug., 1960, Vienna. (Larvae and adults of Trogoderma granarium Everts were found in February 1953 at Ancona, Italy, in wheat imported from Turkey. Further inspections in 1954-59 showed that the beetle was present in cargoes from various countries of the Middle East, and local infestations were found to have become established at Trapani (Sicily), Gravina and Castellammare di Stabia. Control and preventive measures were intensified, and the beetle is thought not to be permanently established anywhere in the country, although the climate in many areas seems suitable. A related species imported in wheat from North America was identified as T. inclusum Lec.)

Darling, H.S., 1951. Insects and grain storage in the Sudan. Sudan Notes and Records 32: 131-149. (Trogoderma granarium is a pest of sorghum in Sudan)

Day, Cheryl and White, Ben. 2016. Khapra beetle, Trogoderma granarium interceptions and eradications in Australia and around the world. School of Agricultural and Resource Economics (SARE), University of Western Australia, Crawley, Australia Working paper 1609. In early 2013 live khapra beetle larvae were found by Customs and Border Protection at Pembina, North Dakota in a seemingly low-risk shipment of clothes moving from Alberta to Texas. In 2014, larvae and cast skins were detected in dried pigeon peas in passenger luggage arriving from Sudan; in a traveller’s rice package from Saudi Arabia; and stopped in Abu Dhabi, United Arab Emirates in dried chick peas destined for the U.S. In five cases in Dallas, passengers had not declared packages of rice, dried beans, coriander seeds and fava beans from Sudan and India that carried cast skins and live larvae. At the Lewiston Bridge on the Canada and U.S. border, three live larvae were detected in a sea container being hauled into the U.S carrying a shipment of rain ponchos from China. Organic soybeans from India were intercepted in November 2014, and split lentils and spices from United Arab Emirates were intercepted in January 2015. Other live intercepts in 2015 include, declared rice from Saudi Arabia via London; a 2 kg bag of dried beans originating from Somalia; undeclared dried chick peas and peanuts originating from India; and a 13.7 tonne shipment of chickpeas, lentils and other Indian foodstuffs refused entry and re-exported. Cast skins were also detected on dried hibiscus leaves that were packaged with other plastic bags of rice and spices from Iraq. In early 2016, a 25 tonne shipment of cumin seed from India infested with dead larvae was sealed and re-exported.

Douglas, J. W. 1860. (Occurrence of Trogoderma sp.) Transactions of the Royal Entomological Society of London 5: 113. (Record of occurrence of Trogoderma granarium in London as early as 1860 in rice from Burma.)

Durrant, John Hartley 1921 Insects associated with grain, etc. Rept. Grain Pests (War) Committee, R. Sot. (London). 9: 33-52. (Introduced in England with consignment of barley from India.)

Eliopoulos, P. A. 2013. New approaches for tackling the khapra beetle.CAB Reviews 8(12): 1-13. (Reviews published research on pest management of khapra beetle.)

Emery, R.N. 1999. Warehouse beetle (Trogoderma variabile). Farmnote No. 77/99. Department of Agriculture, Western Australia.

Emery, R.N., E. Ernestos Kostas, M. Chami. 2008. An urban eradication of khapra beetle in Western Australia. In: D.L. Guo, S. Navarro, Y. Jian, T. Cheng, J. Zuxun, L. Yue, L. Yang, W. Haipeng (Eds.), Proceedings of the 8th Interna­tional Conference on Controlled Atmospheres and Fumi­gation in Stored Products, 21-26 Sept. 2008, Chengdu, China, Sichuan Publishing House of Science and Technol­ogy, Chengdu, China.

Emery, R. N., M. Chami, N. Garel, E. Kostas, and D. C. Hardie. 2010. The use of hand-held computers (PDAs) to audit and validate eradication of a postborder detection of khapra beetle in Western Australia, pp. 1031-1037. In M. O. Carvalho, P. G. Fields, C. S. Adler, F. H. Arthur, C. G. Athanassiou, J. F. Campbell, F. Fleurat-Lessard, P. W. Flinn, R. J. Hodges, A. A. Isikber, S. Navarro, R. T. Noyes, J. Riudavets, K. K. Sinha, G. R. Thorpe, B. H. Timlick, P. Trematerra and N. D. G. White (eds.), Proceedings of the 10th International Working Conference on Stored Product Protection, 27 June-2 July 2010, Estoril, Portugal. Julius Kühn-Institut, Berlin, Germany.

Emery, R. N., M. K. Nayak, and J. C. Holloway. 2011. Lessons learned from phosphine resistance monitoring in Australia. Stewart Postharvest Review doi: 10.2212/spr.2011.3.6 ed.

European and Mediterranean Plant Protection Organization (EPPO). 2002. Diagnostic protocols for regulated pests, Trogoderma granarium. OEPP/EPPO Bulletin 32: 299310.

Freeman, J. A. 1951. Pest infestation control in breweries and maltings. Journal of the Institute of Brewing 57(5): 326-337. Risk of introduction of khapra beetle is at a minimum, since foreign barleys are not used; hence there is every opportunity for the eradication of these pests from maltings without fear of re-introduction.

Freeman, J. A. 1958a. Infestation of stored products in Iran, Report of a survey carried out in October, November 1957 and January, 1958. Infestation Control Laboratory, Ministry of Agriculture, Fisheries and Food and Central Treaty Organization, London. 84 p.

Freeman, J. A. 1958b. Infestation of stored products in Pakistan, Report of a survey carried out in November, December, 1957 and January 1958. Infestation Control Laboratory, Ministry of Agriculture, Fisheries and Food and Central Treaty Organization, London. 130 p.

Freeman. J. A. 1962. Stored products infestation and its control in Turkey, Report on a visit to Turkey in September, 1957. Infestation Control Laboratory, Ministry of Agriculture, Fisheries and Food and Central Treaty Organization, London, 86 p.

Freeman, J. 1976. Problems of Stored Products Entomology in Britain Arising Out of Import of Tropical Products. Ann. Appl. Biol. 84:120-124. (Today Trogoderma granarium is regularly introduced on oilcakes from Burma, India, Sudan and Nigeria, but was probably introduced to maltings between 1908 (Walker 1917) and 1918 (Mason 1921) in barley from Indian subcontinent. It must have been introduced earlier many times but fail to establish. Douglas (1860) reported khapra beetle in London in rice from Burma. In modern maltings, khapra beetle infestations are prevented by storage of malt under cool conditions.)

Freeman, J. A. 1974. A review of changes in the pattern of infestation in international trade. EPPO Bull. 4: 251-273. Trogoderma granarium may be able to establish in heated buildings such as maltings in temperate climates. Khapra beetle was intercepted 46-131 times per year (6-18 times per 1000 cargos inspected) between 1967 and 1973 in rice, rice bran, groundnuts, cotton seed or groundnut seed cake and gum imported into Great Britian from India, Myanmar, Nigeria and Sudan. In 1969, there was an increase in interceptions on groundnut cake from Nigeria but by 1973 incidence was at a low level again. Rice from Burma is more likely than rice carried in dry cargo containers from Australia and US to be infested by khapra beetle which has not yet (1974) been found in rice from Thailand and China. Trogoderma granarium may be found in rice bran from Myanmar and India and this rice bran is a frequent source of cross infestation of other commodities during international transport. Review of rice bran infestations from Myanmar has resulted in significant reductions in the proportion of heavily infested cargoes.

A large quantity of goods can be inspected rapidly and economically in ship hold and the cargo treated in port before distribution but this is not possible with shipping containers. Shipping containers have not yet (1974) been used for countries whose exports may be infested by khapra beetle. Transhipment into containers in European ports of infested cargoes from the Far East and their movement by short sea routes to minor ports in Great Britain may avoid inspection. Similarly, the increase use of roll-on, roll-off lorries from as far away as Iran and Afghanistan is a problem. Because of difficulty of inspection of shipping containers on arrival, it would be highly desirable for all cargoes liable to insect infestation to be treated in countries of origin and for spot checks to be made on arrival. There is a risk that insects will be spread by remaining after the removal of the infested load and contaminate future loads.

French, S. and R.C. Venette. 2005. Mini Risk Assessment, Khapra Beetle, Trogoderma granarium (Everts) (Coleoptera: Dermestidae). USDA–APHIS–PPQ–Cooperative Agriculture Pest Survey–Pest Risk Assessment (PRA).

Froggatt, W. W. 1921. “Khapra”, an Indian wheat pest. Agr. Gaz. N.S. Wales. 32(1) :21-23. Rev. Appl. Ent., ser A, 9: 218. Lefroy said that khapra beetle has been found in wheat in Sidney, Australia but no khapra beetle were found with a very thorough search of the stacks at White Bay and Enfield nor at other locations. There is reason to believe that khapra beetle would not do as much damage as under the dry summer heat of Northern India if it were introduced into Australia where climate is different.

Hammond, N. E. B., D. Hardie, C. E. Hauser, and S. A. Reid. 2016. Can general surveillance detect high priority pests in the Western Australian Grains Industry? Crop Protection 79: 8–14.

Hammond, N. E. B., D. Hardie, C. E. Hauser, and S. A. Reid. 2016. How would high priority pests be reported in the Western Australian grains industry?. Crop Protection 79: 26-33.

Harris, D.L. 2009. Khapra Beetle, Trogoderma granarium Everts (Insecta: Coleoptera: Dermestidae). University of Florida, IFAS Extension (EENY-372 (IN667).

Honey, Sabyan F., Babar Bajwa, Muhammad S. Mazhar and Waqas Wakil. 2017. Trogoderma granarium (Everts) (Coleoptera: Dermestidae), an alarming threat to rice supply chain in Pakistan. International Journal of Entomological Research 5:23-31. (In 2011, Khapra beetle larvae were detected twice in USA in rice shipped from Pakistan and in 2014, 43000 pounds of infested rice was returned to Pakistan. In 2013, 3000 metric tons of rice from Pakistan infested with Khapra beetle was rejected at Mexico border and returned. Khapra beetle in rice and stiff competition in world rice market resulted in decline of rice exports from 42 million tons to 37 million and 1000 million dollar loss to rice industry in Pakistan.)

Howe, R. W. 1952. Entomological problems of food storage in northern Nigeria. Bulletin of Entomological Research 43: 111-144. The khapra beetle was first observed in Nigeria in 1948 on stored sorghum (guinea corn) stored as a reserve for feeding school children and civil prisoners in Kano. Infestation was eradicated but in January 1949 infested groundnut storage was discovered and khapra beetle was soon too widespread to eliminate, being found at Kaura Namoda 140 miles west north-west of Kano. Evidence was obtained that it had been present on groundnuts in 1946 and probably in 1944. Adults cannot fly and both adults and larvae can be dispersed by wind and on the clothing of workers. The larvae feed on the groundnuts, powdering more than they consume, and also cause weakening of the sacks which ultimately tear. Before 1940 groundnuts were all moved out of storage by April of each year, but in 1946 some were still in storage in November when the next crop was harvested and with overlapping crops since then some were stored for 18 months. Petty theft of nuts and sale of these nuts with the new crop increased the khapra beetles around at the beginning of storage.

Hopkins, L. 1955. Food preferences of the khapra beetle. Jour. Econ. Entomol. 48(3): 332-333.

Howe, R., and D. Lindgren. 1957. How Much can the Khapra Beetle Spread in the USA. J. Econ. Entomol. 50: 374-375.

Howe, R.W. 1958. A theoretical evaluation of the potential range and importance of Trogoderma granarium Everts (Coleoptera. Dermestidae) in North America. Proceedings of the tenth International Congress on Entomology 4: 23–28.

Hunter, F.A., Jean B.M. Tulloch and M.G. Lambourne. 1973. Insects and mites of maltings in the East Midlands of England Journal of Stored Products Research 9(2): 119-141. (By 1964, many of the old conventional floor maltings had been replaced by new, more efficient mechanized plant. Few (3 out of 36) old malt stores close to the warmth of the kilns and with wooden bins that had many cracks retaining residues yielded live Trogoderma granarium and none was found in any new store.)

Hurlock, E. T. 1961. Persistence of khapra beetle in ships’ holds. Pest Technology 3: 144-146 (Mar).

Hurlock, E. T. 1962. New efforts at khapra beetle control. Pest Technology 4(7): 150-151 (Apr).

Kenney, M., Orr, D., and Shannon, M. J. 1991. Regulatory and inspection functions in the U.S. Department of Agriculture. Pages 505-518 in: Ecology and Management of Food-Industry Pests. Food and Drug Administration Tech. Bull. 4. J. R. Gorham, Ed. Assoc. Off. Anal. Chem., Arlington, VA. (From 1971 to 1981, khapra beetle was intercepted at ports 1,336 times, 74% in New York and San Francisco. Majority of interceptions were in gum, sheepskins and artware. In 1980, an infestation was discovered in a spice warehouse in New Jersey. Subsequently, 25 infestations were found in six states, i. e., California, Maryland, Michigan, New Jersey, New York, Pennsylvania, and Texas were discovered and eradicated)

Kerr, J. A. 1981. Khapra beetle returns. Pest Control 49(12): 24-25. In 1980, APHIS discovered a khapra beetle infestation from India in a New Jersey spice warehouse and shortly after in Baltimore, Maryland and Michigan and then warehouses in California, Pennsylvania and Texas receiving shipments from infested warehouse. APHIS found 20 infestations and all but one were fumigated.

Khidr, Sahand K., Waran Nooraldeen A. Agha, and Adil H. Amin. 2017. Molecular Identification of Three Stored Product Insect Species on Dried Fruits and Their Control with the Use of Some Aqueous Plant Extracts. Science Journal of University of Zakho 5(2): 178-186.

Kingsolver, J. M. 1963. Notes on the pictorial key for identification of the larvae of dermestid genera. U.S. Dept. Agr. Coop. Econ. Ins. Rpt. 13(15): 385-386.

Klassen, W. 1989. Eradication of introduced arthropod pests: Theory and historical practice. Miscellaneous Publications Entomological Society of America 73: 1-29. (The cost of eradications was $8.4 million spent by federal government and an additional $6.5 million spent by property own­ers.)

Kraszpulski, P. 1985. Khapra bettle – a quarantine pest Trogoderma granarium Everts. Och. Ros. 29: 8-9.

Lepesme, P. 1938. Sur Ie regime et l’importance economique de quelques Trogoderma (Coleoptera Dermestidae). (On the diet and economic importance of some Trogoderma (Coleoptera Dermestidae)) Rev. Franc. d’Ent. 5(2) :104-6. Rev. Appl. Ent., ser. A, 27: 196. (Fourth instar larvae infest sound grains, but younger ones can only attack damaged grain)

Lebedev, V. A. and Saplina, G. S. 1978. The khapra beetle Trogoderma granarium: Quarantine pest of stored products Zashchita rastenii ot vreditelei i boleznei 1: 44. Trogoderma granarium Everts is not established in the USSR but is occasionally found in imported stored materials and quarantine regulations are in force to prevent its introduction. In 1977 it was intercepted in the USSR on polyethylene and cotton yarn from Vietnam.

Lin, C. K. 1971. Pest of stored grain and grain products in Taiwan I. Studies on the biology of the khapra beetle, Trogoderma granarium Everts. (Coleoptera: Dermestidae) (in Chinese). Plant Prot. Bull. 13: 18-24.

Lin, C. K. 1973. Biological studies on the khapra beetle Trogoderma granarium Everts (Coleoptera: Dermestidae) II Studies on the effect of temperature in relation to its larval stadium (in Chinese). Plant Prot. Bull. 15: 86-92.

Lin, T., and C. H. Li. 1983. Studies on ecology of khapra beetle, Trogoderma granarium E. J. Agric. Res. China 32: 383-389.

Lindgren, D., L. Vincent, and H. Krohne. 1954. Khapra beetle in California: Eastern hemisphere insect destructive to stored grain, cereal products and foodstuffs established in state. California Agriculture 8(9): 7,15.

Lindgren, D. L. 1955. Watch for the khapra beetle in stored foods…in barley for example. California Agricultural Experiment Station Leaflet 51

Lindgren, D.L., Vincent L.E., and Krohne H.E. 1955. Khapra Beetle Control Studies: preliminary results if tests with fumigants and dust give promise of effective treatments against destructive pest. California Agriculture, March 1955.

Lindgren, D. L., Vincent, L. E., and Krohne, H. E. 1955. The khapra beetle, Trogoderma granarium Everts. Hilgardia 24:1-36. (Khapra is Indian word for brick describing the habit of beetle aestivating in the pores in bricks.)

Ma, J., Hu, X.N., Liang F., Wang, X.G. and Zhao, J.P., 2009, Molecular identification for Trogoderma granarium Everts and Trogoderma glabrum (Herbst), Zhiwu Jianyi (Plant Quarantine) 23(4): 27-28

Ma, Ping, XiaoLong Jiang, and ZhengYue Li. 2009. Risk analysis of Trogoderma granarium in Yunnan by comprehensive evaluation index system for invasive alien pests in Yunnan. Southwest China Journal of Agricultural Sciences 22(3): 861-864.

Marcovitch. S. 1954. The insect pest situation in Israeli agriculture. J. Econ. Entom. 47: 19-23. (Trogoderma granarium is present in serious numbers in the Jordan Valley and without proper control measures damage may frequently be 30%.)

Martínez-Jacinto, Nicolás G., Ismael Delgadillo Villanueva, Rubén Hernández-Rivero, José Abel, and Rigoberto González-Gómez López-Buenfil. 2017. Estrategias de Detección Para Palomilla Gitana Asiática (Lymantria dispar asiatica) (Lepidoptera: Lymantriidae) y Gorgojo Khapra Trogoderma granarium (Everts) (Coleoptera: Dermestidae). (Detecction strategies for Asian gypsy moth Lymantria dispar asiatica (Linnaeus) (Lepidoptera: Lymantriidae) and khapra beetle Trogoderma granarium (Everts)). Entomología Mexicana 4: 390−395.  Due to the importance of khapra beetle to Mexico, the Servicio Nacional de Sanidad, Inocuidad y Calidad Agroalimentaria (SENASICA), through Phytosanitary Epidemiological Surveillance Program, implemented during 2016 trapping program in the states of Baja California, Campeche, Colima, Chiapas, Chihuahua, Michoacán, Nuevo León, Oaxaca, Sinaloa, Sonora, Tamaulipas, Veracruz, and Yucatán. Trogoderma granarium was not detected in Mexico and is classified as quarantine pests.

Mason, L. 2002. Khapra beetle. Pest not found in the U.S., but a potential threat anyway. Grain Journal 30(4): 34-35.

Mason, Linda 2013. Beetlemania – of another kind, Khapra beetle is back……

Mason, F.A. 1921. The destruction of stored grain by Trogoderma khapra, Arrow. A new pest in Great Britain. Bur. Bio-Technol., Leeds, Bul. 2:27-38. Appl. Ent. Ser. A. 9: 143.

Mawlood, N. A., and M. S. Abdul-Rassoul. 2000. Notes on Trogoderma species (Coleoptera, Dermestidae) of Iraq. Bulletin of the Iraq Natural History Museum 9(2): 51-60.

McElwee, Hamish 2000. The economics of targeted surveillance for khapra beetle in Western Australia: a preliminary analysis. Bulletin (Western Australia. Agriculture Western Australia)

McElwee, H. 2000. The potential economic impact of khapra beetle (Trogoderma granarium  Everts) on the Western Australian wheat industry. Bulletin (Western Australia. Agriculture Western Australia)

Mendenhall, W. 1956. Report on the Khapra Beetle Program in Arizona. J. Econ. Entomol. 49: 508-511. (Between May 28 and November 8, 1956, 30 properties have been treated with eradication type fumigations. Fourteen were farm and ranch properties and 16 were commercial type establishments. Approximately 10,800,000 cubic feet of warehouse space was fumigated out of an estimated 30 million cubic feet which was known to be infested. These properties range in size from a small feed store containing 5,100 cubic feet to a mill and elevators of 2,600,000 cubic feet. The cost of eradication for Arizona was calculated.)

Mitsui, E. 1965. Possibility of the establishment of the khapra beetle in Japan. I. Can the khapra beetle establish in Japan? (in Japanese) Bull. Food Research Institute Tokyo 19: 30-36.

Mordkovich, B., and A. Sokolov. 1992. To eliminate colonies of Khapra beetle (in Russian). Zashch. Rast. 6: 49-52.

Mordkovich, Y. B. and E. A. Sokolov. 1999. The determinant of quarantine and other dangerous pests of raw materials, stock and seed products. Kolos, Moscow 384.

Mordkovich, Ya B., and E. A. Sokolov. 2000. Detection of Khapra beetle in warehouses. Zashchita i Karantin Rasteniĭ 12: 26-27.

Munro, J. W. 1935. The khapra beetle. Journal of the Institute of Brewing 41: 373-374. (First recorded in England in 1917 when Arrow again described it as a new species.)

Munro, J.W. 1940. Report on a survey of the infestation of grain by insects. London Dep. Sci. Industr. Res. 54 pp. Warmth of kiln and malt stores are essential to the development and population growth of Trogoderma granarium and this species is of greatest importance to the brewing and distilling industry in England. Dead larvae render grain sour and raise the water content of malt, making it unfit for use by brewer. The use of English barley is increasing but some is still imported from California, Poland, Australia, India and Chile. Thirteen out of 27 maltsters refuse khapra beetle infested grain and one grain merchant refused delivery of Indian barley to his customer when he found it infested in the ship. In spite of its know preference for warm temperatures, it was found in large numbers in groundnut cake and Indian wheat cargoes on ships and in the fabric of a warehouse in the port of Aberdeen during March. Brewers, maltsters and distillers have spent large sums in replacing wooden bins with steel bins, making other structural alterations and fumigating to eliminate khapra beetle. Complete demolition of two malt bins and treatment of the walls with a blowlamp were successful in its eradication at a distillery in the Highlands. Wet and dried grain, malt culmings and spent hops by-products sent directly from malting to agricultural merchants and farms are occasionally infested by Trogoderma.

Mookherjee, P. B., M. G. Jotwani, P. Sircar and T. D. Yadav. 1968. Studies on the incidence and extent of damage to insect pests in stored seeds. I. Cereal seeds. Indian J. Entomol. 30: 61-65. Trogoderma granarium was found infesting wheat in 4 of 7 geographical areas of India and was not found in southern, central and eastern areas. It was found on barley in northern region. The lowest percentage damage in the southern area may be a result of Trogoderma granarium being absent. Humidity was lower in areas where khapra beetle was present.

Mookherjee, P. B., M. G. Jotwani, T. D. Yadav, and P. Sircar. 1970. Studies on incidence and extent of damage due to insect pests in stored seeds. II. Leguminous and vegetable seeds. Indian J. Entomol. 32: 350-355. (Trogoderma granarium was found on 4 of 26 vegetable seeds in central west (3), central east (1) and northern (1) areas of India.)

Morison, G. D. 1925. The khapra beetle (Trogoderma granarium Everts). Proc. R. Phys. Soc. Edinb. 21: 10-13.

Myers, S. W. and D. W. Hagstrum. 2012. Quarantine. p. 297-304. In D. W. Hagstrum, T. W. Phillips and G. Cuperus (eds) Stored Product Protection S156. Kansas State University, Manhattan, KS

Nakayama, Shonosuke. 1932. On the ecology, and especially the moults, of Trogoderma granarium Everts. Oyo-Dobutsugaku Zasshi. (Jap. Soc. Appl. Zool.) 4(3): 150. Rev. Appl. Ent., ser. A, 20: 605. (Trogoderma granarium found in Korea and Japan, where it was feeding on stored rice and wheat.)

Nguyen, C., D. Lovell, R. Oberprieler, D. Jennings, M. Adcock, E. Gates-Stuart, and J. La Salle. 2013. Virtual 3D models of insects for accelerated quarantine control , pp. 161-167. In Anonymous Institute of Electrical and Electronics Engineers (IEEE) International Conference on Computer Vision Workshops.

Noon, Zenas Barnard. 1957. Food habits and the feeding mechanism of the khapra beetle larva, Trogoderma granarium Everts. MS thesis, University of Arizona.

Noon, Z. B. 1958. Food habits of the khapra beetle larva. J. Econ. Entomol. 51: 465-467.(Larvae prefer vegetable foods, but feed on substances of animal origin. First instar larvae can attack sound grain.)

Nutting, W. L. 1984. History: The khapra story, p. 70–74. Forage Grain: A College of Agriculture Report, University of Arizona, Tucson, AZ.

Nutting, W. L., and Gerhardt, P. D. 1964. A study of the Khapra beetle, Trogoderma granarium, in commercial grain storages in southern Arizona. J. Econ. Entomol. 57:305-314. (Ecological study of the khapra beetle was conducted in Phoenix, Arizona, from September 1955 to January 1956 in commercial warehouse containing about 2500 tons of barley and in an adjacent storage holding about 1300 tons of barley and 1600 tons of sorghum.)

O’Farrell, A. F. and Butler, P. M. 1948. Insects and mites associated with the storage and manufacture of foodstuffs in Northern Ireland. Economic Proceedings of the Royal Dublin Society 3(22): 343-407. (Trogoderma granarium occurred in a distillery silo which had previously contained heated grain. The larvae were living in crevices between the slats of wooden bins; they survived for about a year but eventually died out. The species was also found in a silo attached to one of the Belfast flour mills and dead specimens were obtained from a provender mill in Belfast and a feeding stuffs store in Londonderry.)

Okumura, George T. 1955. Illustrated Key to species of Trogoderma and to related generan of Dermestidae commonly encountered in stored grain in California. Cooperative Economic Insect Report 5(15): 1-5

Okumura, George T. 1966. A supplemental character for separating adult Trogoderma granarium Everts, khapra beetle, from other Nearctic Trogoderma species. Cooperative Economic Insect Report 16(9): 147.

Okumura, G. T. and F. L. Blanc. 1955. Exhibit No. 23. Key to species of Trogoderma and to related genera of Dermestidae. p. 87-89. In Anonymous The Khapra Beetle Trogoderma granarium. Special Report of the Joint Interim Committee on Agriculture and Livestock Problems; Legislature of the State of California. (Adults and mature larvae)

Oliveira, Maria Regina Vilarinho de, S. F. Silva, and K. R. Vilarinho. 2005. Subsídios ao processo de elaboração de plano de contingência: Trogoderma granarium. (Subsidies contingency plan preparation process: Trogoderma granarium) . Embrapa Recursos Genéticos e Biotecnologia. Documentos 135.

Olson, R.L.O., Farris, R.E., Barr, N.B., Cognato, A.I. 2014. Molecular identification of Trogoderma granarium (Coleoptera: Dermestidae) using the 16s gene. J. Pest Sci. 87: 701-710.

Ostrauskas H. and Taluntytė L. 2002. Insects in traps for Trogoderma granarium Everts in Lithuania and warehouse insects in import, export and reexport store products. p.  88-90. In Proceedings of the scientifics international conference Plant protection in the Baltic region in the context of integration to EU, September 26-27, 2002, Kaunas: Akademija

Lithuanian company warehouses (38) were investigated using pheromone traps for Trogoderma granarium and 518 samples from warehouse import, export and reexport products were tested during 1997-1999. Trogoderma granarium was not found or intercepted in Lithuania.

Özberk, Fethiye, İrfan Özberk, Abuzer Yücel, Ayhan Atli, and Duygu İzol. 2017. Khapra beetle (Trogoderma granarium Everts, 1898) in durum wheat (Triticum durum Desf): Impacts on some seed characteristics and marketing price. Türkiye Entomoloji Dergisi 41(2): 207-218.

Padget, L. J. 1954. Status of the Khapra beetle in Western United States Spring 1954. USDA Coop. Econ. Ins. Rpt. 4(28): 557-562. (In 1946, khapra beetle was found at J. B. Hill Warehouse Company, Fresno, California and incorrectly identified by pest control company as black carpet beetle. Fumigation was unsuccessful so by 1949 the 300 tons of grain in warehouse was a total loss and warehouse was no longer used for grain storage. Warehouse was gone over with a blowtorch and residual spray, but used bags were moved to Alpaugh and Angola in Tulare, Co., spreading the infestation, filled with grain and used as bulkheads between piles of loose grain. Two infestations in Portales and Texico, New Mexico can be attributed to shipments of sorghum seed for planting from infested facility in Arizona. Three shipments from same facility to Texas and another to Oklahoma were negative.

In November 20, 1953 Cooperative Economic Insect Report discovery of khapra beetle in California was reported and in December 18 infestation in Phoenix warehouse was reported. R. T. Cotton and K. S. Rohwer were assigned by USDA to assist in survey beginning early April. State Pest Control officials provided valuable assistance in selecting mills to inspect and routes to follow. Nevada, Utah and Oregon supplied personnel for survey in those states. Large wholesale grain-handling establishments were included in survey and also feed-mixing plants because they have a wider assortment of ingredients. For example, Farmer’s Cooperative warehouse had several hundred tons of barley and wheat of local origin. Poultry or Dairy Producers Association had smaller tonnage of grain, but also cotton seed meal from Arizona or California, soybean from Minnesota, corn from Iowa and copra from Philippines.

Feed grains, barley and wheat from Arizona and California generally are not shipped north or east. Negligible amounts are shipped to Nevada, Utah and Colorado. Cottonseed meal and cake for areas west of Rockies is supplied exclusively by Arizona and Colorado. Arizona Certified Sorghum seed is prized throughout western states. Burlap bags are supplied to survey area from Los Angeles, San Francisco and Portland.

All commercial warehoused in Arizona and 150 warehouse in California had been inspected so only a few warehouses in these states were included in the survey. Survey included 67 warehouses in Colorado, 77 in Utah, 29 in Idaho, 21 in New Mexico, 6 in Texas, 10 in Nevada and 36 in Oregon and no khapra beetles were found. No warehouses in Wyoming and Montana were inspected.)

Paini, D. R., and D. Yemshanov. 2012. Modelling the arrival of invasive organisms via the international marine shipping network: a Khapra beetle study 7(9): PloS one e44589  A trace back of the container, which carried the family’s belongings revealed that the container had visited the following ports (in reverse order), Fremantle (Australia), Grangemouth (Scotland), Felixstowe (England), Pt Qasim (Pakistan), Gwangyang (South Korea), Busan (South Korea), Hamburg (Germany), Bangkok (Thailand) (personal communication – Rob Emery, Dept of Agriculture and Food, Western Australia, 2011).

Parker, T. 1922. The suppression of insect pests and fungoid diseases. I. The fumigation of malthouses. Bur. Bio-Technol., Leeds, Bul. 7 :229-34. Rev. Appl. Ent., sere A, 11 :34-35. (Malt stored in cold bins is never affected by the khapra beetle, infestations only appearing where the temperature approximates 90o to 110o F. Generally speaking the greatest infestation occurs around the walls adjoining the kiln or kiln shafts. The larvae migrate from one part of the building to another, but so far have not been observed on barley floors. Fumigation of the whole premises is the best remedy, though it is difficult to remove all the malt and leave the premises vacant for two or three weeks.)

Parker, T. and A. W. Long. 1921. A laboratory note on the control of Trogoderma khapra. Bureau of Bio-Technology Bulletin 4:102-104. RAE A 10: 32 (Khapra beetle is gradually spreading in England and can only be exterminated from malthouses by repeated fumigation.)

Pasek, J. E. 1998. Khapra beetle (Trogoderma granarium Everts): pest-initiated pest risk assessment, USDA APHIS, Raleigh, NC. Khapra beetle infestation was detected in 1997 in a spice processing warehouse in Owings Mills, Maryland. Actions taken to contain and eliminate the infestation included: fumigating all shipments leaving the facility, surface spraying the facility with malathion, vacuuming all surfaces to remove dust and insects, sealing surfaces with several coats of thick latex paint, and decontaminating and replacing spice processing equipment. The methods used likely are not economically feasible for most commodity handlers and were chosen, in part, in this instance because the plant intended to remodel the facility anyway. The facility itself was not fumigated due to citizen opposition.

Six countries (India 34% % > Saudi Arabia 22% > Iran 14% > Pakistan 9% > Kuwait 3% > Sudan 2%) accounted for 84% of interceptions of khapra beetle in US from 1985-1998. During 1971-1984, the order was India 44% > Sudan 24% > Pakistan 15% > Iran 5% > Saudi Arabia = Kuwait < 2%.

Pattinson, I. 1958. Insect infestation in Edinburgh area maltings. Departmental report. Dept. Agric. Fish. Scotland.

Popham, W. L. 1955. Domestic quarantine notices (Plant Pest Control No. 612). Title 7 Agriculture Chapter III Part 301 Subpart khapra beetle, U.S. Department of Agriculture (Lists warehouses, mills and other premises in which khapra beetle detected and designated as regulated areas.)

Qureshi, A. H. 1966. A new approach to eradicate Trogoderma granarium Everts in Kaura Namoda grain stores. Technical Report. Nigerian Stored Products Research Institute, 1965 p. 43-47. In 1965, native storehouses containing mainly beans and grains at Kaura Namoda, Northern Region, Nigeria were heavily infested by Trogoderma granarium. Infestation was eradicated by spraying walls with malathion and fumigating, but a storehouse that had just received a fresh stock of grains was found to be heavily infested by Trogoderma granarium. Continued monitoring will be needed since this has long been a source of infestation from which Trogoderma granarium can spread southward along the trade routes.

Rajapakse, Rohan and Wolly Wijayaratne. 2017. Khapra Beetle and Future of Sri Lankan Tea Export.

Rebolledo, R. and M. Arroyo 1993. Prospección de Trogoderma granarium Everts (Coleoptera: Dermestidae) mediante trampas de feromonas en Madrid. (Prospecting for Trogoderma granarium Everts (Coleoptera: Dermestidae) through pheromone traps in Madrid) Boletin de Sanidad Vegetal. Plagas (Espana) 19: 361-367. (Pheromone trap in Madrid, Spain attracted Trogoderma inclusum but not Trogoderma granarium)

Rebolledo, R. & Arroyo. M., 1994. Prospection of species of Trogoderma (Coleóptera: Dermestídae) by means of traps of pheromone in Madrid, second year of observations. Bol. San. Veg. Plagas, 20(1): 49-56. (During second year, pheromone trap in Madrid, Spain attracted Trogoderma inclusum but not Trogoderma granarium)

Reddy, D. B. 1969. Distribution of khapra beetle in the South East Asia and Pacific region. FAO PIant Protection Committee for S.E. Asia and Pacific Region. Information Letter No. 70.

Rees, D.P. and Banks, H.J. 1999. The khapra beetle, Trogoderma granarium Everts (Coleoptera: Dermestidae), a quarantine pest of stored products: review of biology, distribution, monitoring and control. Canberra, CSIRO Stored Grain Research Laboratory.

Ring, W. M. 1965 Report to the Government of Turkey on a Survey of Khapra Beetle Infestation in Stored Products. Report/Expanded program of technical assistance FAO 2038 (map).

Risbec, J. and Mallamaire, A. 1949. Les animaux predateurs et les insectes parasites du riz cultive en Afrique occidentale. Agron. Trop. Nogent. 4: 70-76. (Trogoderma granarium pest of stored rice.)

Salmond, K. F. 1958. Studies on Trogoderma granarium Everts (Dermestidae-Coleoptera). I. Its importance as a pest of stored maize in the Federation of Rhodesia and Nyasaland, pp. 35-49. In Anonymous Stored food products/Les produits alimentaires emmagasinés. Meeting of specialists on stored food products. Commission for Technical Co-operation in Africa South of the Sahara Publication No. 31, 1957, Salisbury.

Salmond, K. F. 1958. Studies on Trogoderma granarium Everts (Dermestidae-Coleoptera): II. Numbers, distribution and activity of Trogoderma granarium in the fabric of a Cape-steel storage shed, pp. 51-57. In Anonymous Stored Food Products/ Les produits alimentaires emmagasinés. Meeting of specialists on stored food products. Commission for Technical Co-operation in Africa South of the Sahara Publication No. 31, 1957, Salisbury. Wind currents tended to carry smaller khapra beetle larvae from the walls of shed, some larvae to heights of 12-15 ft above ground. Trogoderma granarium larval (adult) populations on girders were 1915 (100) on rail side of shed; 2584 (570) on south end; 54 (8) on other ground level girders; 166 (82) 5 ft above ground level girders;  8 (nil) and 124 (311) on top girders. Totals of 3150 larvae and 20 adults were collected per 40 sq ft of brick and concrete wall base; 2836 larvae and 45 adults were collected per 40 sq ft of floor/wall edge; 1217 larvae and 73 adults were collected per 100 sq ft of floor area below edges of ruberoid felt covering concrete; 174-291 larvae and 19 to 53 adults were collected per 40 sq ft of wall. Most of larvae were alive; most of adults dead and fewer were collected from outside surfaces of the warehouse. Ground sample and maize spillage 6 ft from warehouse had nil khapra beetle larvae and adults. These residual populations in warehouse make regular sweeping of floor and cleaning girders important before insecticide application.

Shannon, M.J. 1989. Chapter 7: Khapra beetle, pp. 109-126. In: Kahn, Robert P., ed. Plant Protection and Quarantine, Vol. II: Selected Pests and Pathogens of Quarantine Significance. CRC Press, Boca Raton, FL.

Between 1971 and 1984, the frequency in percent of khapra beetle interception in United States were by commodity: Gum (17.4%), Sheepskin (16.0%), Artware (13.2%), Capsicum (10.7%), Cotton piece goods (6.2%), Cucurbit (5.4%), Cumin (5.3%), Carpets (4.4%), Foodstuff (3.6%), Automobile (3.2%), Celery seed (2.2%), Cassia (2.1%), Brassware (2.0%), Hibiscus (1.3%), Peanut (1.1%), Rice (1.1%), Pistachio (1.1%), Screens (0.9%), Coffee (0.8%), Niger (0.7%), Goatskins (0.7%), Container (0.7%) with a total of 757 interceptions. Number of other identified hosts with less than five interceptions was 44, total identified hosts = 801 (60%) and total unidentified hosts was 535 (40%). Between 1971-1984, frequency in percent of khapra beetle interceptions in imports into United States by country was India 44% > Sudan 24% > Pakistan 15% > Iran 5% > Saudi Arabia 2%.

For imports from Pakistan into United States for 1982 through 1984, frequency in percent of interceptions of khapra beetle by commodity were Cumin seed (43.2%), Curcurbit seed (18.9%), Bagging (with cargo) (10.8%), Foodstuffs (10.8%), Rice (bagging) (2.7%), Rosa sp. (2.7%), Chickpea (2.7%), Fennel (bagging) (2.7%), Sheepskins (2.7%), Cargo not identified (2.7%) with a total of 37 interceptions

 For imports from India into United States for 1982 through 1984, frequency in percent of interceptions of khapra beetle by commodity were Crating (26.2), Cargo not identified (18%), Celery seed (bagging) (9.0%), Gum (bagging) (8.2%), Gum (8.2%), Foodstuffs (bagging) (5.7%), Brassware (crating) (5.7%), Cumin seed (bagging) (4.1%), Rice (bagging) (2.5%), Niger seed (bagging) (1.6%), Cloth (bagging) (1.6%), Dill seed (1.6%), Coriander (bagging) (1.6%), Coffee (bagging) (0.8%), Lentil (bagging) (0.8%), Carpets (0.8%), Beans (bagging) (0.8%), Psyllium sd. (0.8%), Sheepskins (0.8%), Bagging as cargo (0.8%) with a total 122 interceptions.

For imports from Sudan into United States for 1982 through 1984, frequency in percent of interceptions of khapra beetle by commodity were Cucurbit seed (26.6%), Sheepskins (25.0%), Cargo not identified (17.2%), Hibiscus (bagging) (14.1%), Cassia senna (bagging) (7.8%), Gum (bagging) (6.3%), Pepper (bagging) (3.1%) with a total 64 interceptions.

 The frequency in percent of the pathways of khapra beetle entry into United States for 1982 through 1984 were Cargo (56.7%), Baggage (29.9%), Ship’s stores (9.5%), Holds (3.2%), Mail (0.5%), Ship’s quarters (0.2%) with a total of 432 interceptions of khapra beetle.

Most khapra beetle infested commodities were accompanied by burlap (bagging). Port of New York accounts for 65% of all cargo interceptions of khapra beetle and eight ports account for 95% of interceptions. Interceptions associated with crating, usually a single specimen, only involved India during 1984 and 1985. There was a recent upsurge of infested celery seeds. Threat posed by carriers to United States has diminished in recent years with the introduction of sea containers and improved sanitation on arriving vessels. In 1979, 33-50% of annual interceptions of khapra beetle were arriving ships and with sea containers they constitute only 9.5% between 1982 and 1984.

Shapas, T. J.; Burkholder, W. E. 1979. Program for using sex pheromones to detect khapra beetle at U.S. ports. Proceedings of the North Central Branch of the Entomological Society of America  33: 30. The trapping method used in a survey and detection programme against Trogoderma granarium Everts at ports in the USA is described. The traps contain dichlorvos (DDVP) for killing the insects, wheat-germ oil to attract the larvae, and 1 mg of the sex pheromone of the species ((Z)-14-methyl-8-hexadecenal) to attract adult males. The use of such traps should greatly increase sensitivity for detection of the pest.

Shea, G., J.H. Botha and R.N. Emery 2000. Khapra beetle; Trogoderma granarium. An exotic threat to Western Australia. Factsheet No. 22/2000. Department of Agriculture, Western Australia.

Shen Z.P. 1994. Immunological characterization of Trogoderma granarium, Liangshi Chucang (Grain Storage) 6: 44.

Shepherd, D.R. 1957. Khapra beetle eradication. FAO Plant. Prot. Bull. 5: 75-77.

Sinha, R. N. 1963. Suitability of climatic areas of Canada for infestation of some major stored grain insects on farms. Proceedings of the Entomological Society of Manitoba 19: 31-39.

Sinha. R. N. and Utida. S. 1967. Climatic areas potentially vulnerable to stored product insects in Japan. Appl. Em. Zool. 2. 124-132.

Smetnik, A. I., and L. M. Nikritin. 1986. The khapra beetle. Zashchita Rasteniĭ (Moskva) 10: 41-43.

Smit, Bernard. 1957. The control of stored grain insects in South Africa. Bull. Dept. Agric. S. Afi. No. 355. 36 pp. (Trogoderma granarium was first found in maltings in a brewery in Pietermaritzburg in 1954 and much more serious outbreak in a large mill near Pretoria in 1956. In mill, an accumulation of sweepings and tailings from cleaning machinery had been piled up in bags for ca. 3 years and some had been sold without record, so khapra beetle is likely to be widely distributed.)

Smith, K. G. 1963. Study of an insect population living on bagged groundnuts stored in southern Nigeria with particular reference to the behavior of Trogoderma granarium Everts (Col., Dermestidae). Journal of the West African Science Association 8: 44-57.(Trogoderma granarium population in an imported 24 bag stack were monitored for 18 months to determine whether they might become established in southern Nigeria as a result of introduction on northern Nigerian groundnuts. Despite humid conditions of southern Nigeria, even in small stack, local hot dry conditions which developed near the center were favorable for khapra beetle development. Although khapra beetle infested groundnuts from northern Nigeria have been shipped through the ports of southern Nigeria it does not yet appear to be established there.)

Sokolov, Y.А. 2006 Analysis of phytosanitary risk of khapra beetle Trogoderma granarium for the territory of the Russian Federation. Federal Service for Veterinary and Phytosanitary Supervision, Federal State Institution, All-Russian Center of Plant Quarantine, Moscow.

First khapra beetle infestation in the territory of former USSR was found in 1986 in Turkmenistan in malt at Merken brewing plant. In the next two years, 21 infestations were found (Turkmenistan (8), Uzbekistan (4), Tajikistan (5), Russia (2) and Kazakhstan (2)) and infestations in Turkmenistan were eradicated within 3 years. The first detection of khapra beetle in warehouses in the territory of Russia was in May 1987 and two additional infestations were detected in the Stavropol region with live larvae in the batch of fodder corn imported from the United States to the Kochubeevskiy plant of cereal products and Nezlobnenskiy feed milling plant. Both were eradicated.

Sonda, M. 1968. The status of Trogoderma granarium Everts and T. varium (Matsumura and Yokoyama) (Col., Dermestidae) as pests of stored products in Japan. J. Stored Prod. Res. 4: 23-30. (Khapra beetle was first reported in Japan in 1923 and was absent by 1950 when the law to inspect imported stored products was introduced. In 1964, a nation-wide search of 30 properties in Japan found khapra beetle at four breweries some distance from each other in Tokyo, Osaka, Hyogo Prefecture and Fukuoka Prefecture and in two maltings in Tochigi Prefecture near Tokyo and in Shiga Prefecture near Osaka that supplied these breweries. An eradication program involving fumigation with methyl bromide at infested premises, and filling crevices of warehouses and silos with mortar was undertaken.)

Stanford, C. L. 1956 The khapra beetle. Cereal Science Today 1: 134-135

Stibick, J. 2007. New Pest Response Guidelines: Khapra Beetle. USDA–APHIS–PPQ–Emergency and Domestic Programs, Riverdale,Maryland.

Strong, R. G. and R. G. Aundt. 1962. Crossbreeding Studies with Seven Species of Trogoderma. J. Econ. Entomol. 55(4): 445-448. (Reproductive isolation was complete for five species, namely, T. grassmani, T. inclusum, T. parabile, T. simplex, and T. sternale. Reproductive isolation between T. granarius and T. glabrum was not complete, as interspecific hybrid offspring were readily obtained from crosses between these two species, but the hybrids were sterile.)

Strong, Rudolph G., and George Okumura. 1966. Trogoderma species found in California. Bulletin Department Agriculture, State of California 55(1): 23–30.

Strong, R. G., Okumura, G. T., and Sbur, D. E. 1959. Distribution and host range of eight species of Trogoderma in California. J. Econ. Entomol. 52:830-836. (Distributions of eight species of Trogoderma (glabrum, granarium, grassmani, inclusum, ornatum, variabile, simplex and sternale) were mapped. Of 132 host materials found during survey,  glabrum was found in association with 6, granarium with 37, grassmani with 13, inclusum with 33, ornatum with 20, variabile with 77, simplex with 60, and sternale with 79.)

Stuart, Melissa K; Alan V Barak; Wendell E Burkholder. 1994. Immunological identification of Trogoderma granarium Everts (Coleoptera: Dermestidae).Journal of Stored Products Research 30 (1): 9-16

Swaine, G. and Mutter, S. 1961. The khapra beetle. Tanganyika Ministry of Agriculture Bulletin No. 7. Tanzania, Kenya, Uganda and Zanzibar passed legislation requiring compulsory treatment of grains, flours, edible oil seed, pulses and beans from India, Pakistan, Myranmar and Thailand in 1956 when khapra beetle infested spices were imported into Zanzibar from the Far East. Bags from all countries except Kenya and Uganda also required fumigation. Early in 1960, serious outbreaks of khapra beetle were discovered at 8 stores in Dodoma, Tanzania and controlled by fumigating the infested articles and spraying interior of the building with insecticides. Only one store at Dodoma had a second outbreak. Legislation was amended to include additional importing countries and commodities.  Inspection services were expanded beyond port and search revealed a number of very small outbreaks at widely scattered points that were controlled.

Taylor, H. S. 1924. The Khapra beetle: its influence on the moisture content of malt in store. Bur. Bio-Technol. Bul., Leeds 2(13): 113-117. Rev. Appl. Ent., ser. A, 13: 90-91.

Taylor, H. S. 1936. The khapra beetle . Journal of the Institute of Brewing 42: 299. (Warming stored malt is generally the problem and the best solution is to redesign facility to reduce temperature of malt during storage. Heat from kiln or sun shining on the roof can heat up the malt store and insect thrive.)

Tebb, G. 1968 A survey of infestations in maltings and breweries. Journal of the Institute of Brewing 74: 207-219. (Khapra beetle was reported three times as often (3 of 45 malt stores) as in earlier survey by Hunter et al. 1973. Since earlier survey involved inspection and this survey was questionnaire only, much of difference may have been due to misidentification. This is the 5th in a series surveys between 1940 and 1968, i.e., Munro 1940, Freeman 1951, Pattinson 1958, Hunter et al 1973)

Tezcan, Serdar, Yusuf Karsavuran, Esat Pehlivan and Jiri Hava. 2004. Contribution to the Dermestidae (Coleoptera) fauna of Turkey along with new records. Turkish Journal of Entomology 28(1): 27-37. (12 citations for distribution of Trogoderma granarium in Turkey)

Thompson, Michael, Anita Lyons, Lalith Kumarasinghe, Darren R. Peck, Gary Kong, Steve Shattuck, and John La Salle. 2011. Remote microscopy: a success story in Australian and New Zealand plant biosecurity. Austral Entomology 50(1): 1-6.

Vayssiere, P. and Lepesme, P. 1938. Un dangereux parasite des grains emmagasints. Trogoderma granarium Everts. (A dangerous parasite of stored grains. Trogoderma granarium Everts) Agron. colon. 243: 65-71. (With increasingly wide distribution of Trogoderma granarium as a pest of stored grain and the danger of its establishment in France and French colonies, its synonymy, geographical distribution and control are discussed.)

Viljoen, J. 1990. The Occurrence of Trogoderma (Coleoptera, Dermestidae) and Related Species in Southern Africa with Special Reference to T. granarium and its Potential to Become Established. J. Stored Prod. Res. 26: 43-51. The first record of Trogoderma granarium in South Africa was at Pietermaritzburg in May 1954 in a consignment of malt imported from England in 1953. The malt was promptly processed into beer and no more Trogoderma were found in the premises. There is no evidence of spread to other areas. The second, in June 1955, was a heavy infestation in a mill at Bon Accord near Pretoria, in a stack of sweepings and tailings collected from various sources over 3 years. These were promptly milled as stock feed and the area previously occupied by the stack was heavily sprayed with insecticide. Although the infestation had ample opportunity to spread throughout the warehouse and mill and to disperse further, there have been no further records in the Pretoria area. Between 1955 and 1968 there were poorly documented reports of light infestations of “khapra beetle” at Heidelberg, Transvaal (1955) Lichtenburg horizontal bulk storage (1956) and Standerton (1961) where no beetles seem to have been collected, and from Potchefstroom (1964) Klerksdorp (1965) and Pietersburg (1968). The insects from Potchefstroom and Klerksdorp were identified as Trogoderma versicolor (Creutzer) (now inclusum Le Conte), and those from Pietersburg deposited in the National Collection of Insects have been identified as Phradunoma sp. (Jonsson, personal communication). Thus there is no evidence that Trogoderma granarium was found over this period. In 1972 a severe infestation of T. granarium was discovered in locally produced wheat at Onseepkans, a small irrigation settlement some 190 km west of Upington on the south bank of the Orange River.

Voelkel, H. 1924. Zur Biologie und Beklmpfung des Khapraklfers, Trogoderma granarium Everts. (On biology and control of the Khapra beetle, Trogoderma granarium Everts). Arb. biol. Reichsanst. Land-u. Forstw. 13:129-171. (Trogoderma granarium was first reported in Germany in 1921, while it was found in 1923 in large numbers in the malt stores of a brewery in northwestern Germany. Young larva cannot attack whole grain and feed on the floury debris from the feeding of older larvae. Rough cloths suspended over and in contact with grain surface trap Trogoderma granarium larvae prior to pupation and they can be destroyed. Boards, sacks, etc., lying on the surface are hiding places that collect larvae.)

Walker, J. J. 1917. Notes on Trogoderma khapra Arrow, a recently described dermestid granary pest. Entomologists monthly Magazine 53: 165. (Khapra beetle found in screenings of barley from Karachi at Strood, Kent, England on July 30, 1908.)

Ward, A. 1965. The khapra beetle, Trogoderma granarium, and two other species of Trogoderma (Coleoptera: Dermestidae) intercepted entering New Zealand. N. Z. Entomol. 3: 39-41.

Whellan, J. A. 1956. Grain pest new to Rhodesia. Rhodesia Agricultural Journal 53: 41-50. (In February 1955, a severe infestation of the Trogoderma granarium was discovered at Bulawayo, Zimbabwe in beans stored for African rations.)

Whellan, J. A. 1958 Current problems in grain storage in Southern Rhodesia. p. 77-82. In Anonymous Stored food products/Les produits alimentaires emmagasinés. Meeting of specialists on stored food products. Commission for Technical Co-operation in Africa South of the Sahara Publication No. 31, 1957, Salisbury. (Trogoderma granarium Everts was found infesting screenings in Zimbabwe in September 1955, but fumigation was ineffective as a result of leakage. During 1956, Trogoderma granarium was inadvertently distributed on railway trucks and in maize meal or screenings to other locations, where, heavy infestations developed.)

Yulin A, Caihua D, Hongbing Z, Guoyao J. 1999. RAPD assessment of three sibling species of Trogoderma Dejean (Coleoptera: Dermestidae). p. 1755–1757. In: Jin Z, Liang Q, Liang Y, Tan X, Guan L, editors. Proceedings of the 7th International Working Conference on Stored-Product Protection, 14–19 October 1998; Beijing, China. Chengdu, China

Zacher, F. 1922. Eingeschleppte Vorratssehadlingo, (Imported pests of stored products.) Deut. Gesell. f. Angew. Ent. Verhandl. 3. Mitgliederversammlung zu Eisenach 28 bis 30, pp. 55-59. Rev. Appl. Ent., ser A, 11: 130-131. (Pests found in 1921 in grain from foreign sources stored in Germany included the khapra beetle)

Zacher, F. 1926. Der Khaprakafer ein neuer Schadling an Getreide, Malz und Hulsenfruchten. (The Khapra beetle, a new pest of cereals, malt and pulses). Marktber, Hansablum 214, suppmt.: 8-10. Rev. Appl. Ent., ser A, 14: 599.

Zacher, F. 1935. Die Vorratsschädlinge Ägyptens. (The pests of Egypt) Mitteilungen der Gesellschaft für Vorratsschutz 11: 42-44, 55-57, 66-67, 78-82. (Trogoderma granarium was found in grain and malt warehouses of brewery in Egypt. Indicates that khapra beetle is present is India and has been introduced in England, France and Germany. Gives table of the relative percentages of cereals, cereal waste, flour, etc. samples with Sitophilus, Trogoderma and Rhyzopertha at 6 locations and Trogoderma was found in 16.6 to 80% at 4 locations and not found at 2 locations.)

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Zheng Sizhu, Li Jialin, Wei Yihan, Gao Yuan, Zhan Guohui, Yang Xiaojun and Chen Yunfang. 2016. DNA barcoding identification of Dermestidae species, Mitochondrial DNA. Part A, 27(6): 4498-4502.

Zhongping, Xiong, Jiří Háva, and Pan Yongzhi. 2017. A new species of the genus Trogoderma Dejean, 1821 from China (Coleoptera: Dermestidae: Megatomini). Studies and Reports Taxonomical Series 13 (1): 241-247.

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Zungoli, P. 1984. The khapra beetle: serious destroyer of stored grain. Pest Control 52(11): 66.


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