stored-product insects in botanical warehouses

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Stored-product insects in botanical warehouses A.Y. Abdelghany a, b , S.S. Awadalla a , N.F. Abdel-Baky a , H.A. El-Syrafi a , Paul G. Fields b, * a Economic Entomology Department, Faculty of Agriculture, Mansoura University, Mansoura 35516, Egypt b Cereal Research Centre, Agriculture & Agri-Food Canada, 195 Dafoe Road, Winnipeg, Manitoba, R3T 2M9, Canada article info Article history: Received 19 June 2009 Received in revised form 27 October 2009 Accepted 4 November 2009 Keywords: Egypt Lasioderma serricorne Stegobium paniceum medicinal plants abstract Insect pests infesting six stored botanicals were sampled monthly in two Egyptian warehouses over 12 months. The plants sampled were; roselle (Hibiscus sabdariffa), mogat (Glossostemon bruguieri), coriander (Coriandrum sativum), anise (Pimpinella anisum), chamomile (Matricaria chamomilla) and marjoram (Origanum majorana). The warehouses were located in northern Egypt in Mansoura and Bilqas. Lasio- derma serricorne and Stegobium paniceum were the most common insect pests in warehouses. The beetles Tribolium castaneum, Tribolium confusum, Trogoderma granarium and Cryptolestes ferrugineus had lower and similar levels of infestation. The moths, Plodia interpunctella and Sitotroga cerealella, had the lowest infestation levels. There were small differences in infestation by the most common insects, L. serricorne and S. paniceum, among the six botanicals. In general, M. chamomilla and O. majorana had the lowest level of infestation. The other plants, H. sabdariffa and C. sativum had higher levels of infestation. The warehouse in Bilqas had higher populations than the warehouse in Mansoura. In Bilqas, the temperature and relative humidity were slightly higher and the warehouse was older and had open windows, factors that may have contributed to higher insect infestations. Crown Copyright Ó 2010 Published by Elsevier Ltd. All rights reserved. 1. Introduction Most research in stored-product protection focuses on cereals, as cereals are the most common commodity in storage, and there are several insects and moulds that can reduce the quality and quantity of cereals in storage. However, all agricultural products are stored for some time before consumption, and they can also be infested by insects and moulds in storage. Dried plants, roots, stems, leaves, seeds and flowers, have been used from the dawn of history for medicine and as spices for cooking (Craker, 2007). Recently, there has been an increase in the popularity of herbal remedies in North America and Europe. However, there is a growing need to understand the problems in storing botanicals. Botanicals are also attacked in storage by several insects. The most common insects found in botanicals are Lasioderma serri- corne (F.) and Stegobium paniceum (L.) (Lefkovitch, 1967; Tawfik et al., 1984; Awadallah et al., 1990; Arbogast et al., 2002). These insects are even able to develop on Acorus calamus L. rhizomes (Srivastava and Saxena, 1975), a plant that has essential oils that are toxic to several stored-product insects (El-Nahal et al., 1989). There are a number of other insects found in stored botanicals (Araecerus fasciculatus (Degeer), Tenebroides mauritanicus (L.), Tribolium castaneum (Herbst), Oryzaephilus mercator (Fauvel), Ahasverus advena (Waltl), Thaneroclerus buqueti Lefebvre and Thorictodes heydeni (Reitter) (Abraham, 1975; Tawfik et al., 1984; Basak, 1991)). There have been several studies examining insects in stored botanicals worldwide; Egypt (Kamel, 1958; El-Halfawy, 1977), India (Abraham, 1975; Basak, 1991), Germany (Schmidt and Graebner, 1987), Korea (Toh, 1998) and USA (Arbogast et al., 2002). Egypt is a major producer of botanicals. Each year, Egypt exports 7000 to 15,000 t of botanicals (Lange and Mladenova, 1997). These plants are grown in southern Egypt, harvested from June to August, dried, then placed in 50 kg jute bags and stored for 2–10 months in warehouses before being shipped to processing facilities in Egypt, Europe and USA. As in other countries, the two most common insects in botanicals in Egyptian warehouses are L. serricorne and S. paniceum (Tawfik et al., 1984; Awadallah et al., 1990). These and other stored-product insects cause significant damage during storage. To reduce the damage, bag stacks are covered with a tarp and fumigated with phosphine derived from aluminum phosphide (Bond, 1984) once or twice during storage. However, there is an increased demand from customers in Europe and USA for organic production, which does not allow the use of phosphine. Yet there are very low tolerances for insect damage, live insects and insect fragments in stored botanicals. * Corresponding author. Tel.: þ1 204 983 1468; fax: þ1 204 983 4604. E-mail address: paul.fi[email protected] (P.G. Fields). Contents lists available at ScienceDirect Journal of Stored Products Research journal homepage: www.elsevier.com/locate/jspr 0022-474X/$ – see front matter Crown Copyright Ó 2010 Published by Elsevier Ltd. All rights reserved. doi:10.1016/j.jspr.2009.11.001 Journal of Stored Products Research 46 (2010) 93–97

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Journal of Stored Products Research 46 (2010) 93–97

Contents lists avai

Journal of Stored Products Research

journal homepage: www.elsevier .com/locate/ jspr

Stored-product insects in botanical warehouses

A.Y. Abdelghany a,b, S.S. Awadalla a, N.F. Abdel-Baky a, H.A. El-Syrafi a, Paul G. Fields b,*

a Economic Entomology Department, Faculty of Agriculture, Mansoura University, Mansoura 35516, Egyptb Cereal Research Centre, Agriculture & Agri-Food Canada, 195 Dafoe Road, Winnipeg, Manitoba, R3T 2M9, Canada

a r t i c l e i n f o

Article history:Received 19 June 2009Received in revised form27 October 2009Accepted 4 November 2009

Keywords:EgyptLasioderma serricorneStegobium paniceummedicinal plants

* Corresponding author. Tel.: þ1 204 983 1468; faxE-mail address: [email protected] (P.G. Fields).

0022-474X/$ – see front matter Crown Copyright � 2doi:10.1016/j.jspr.2009.11.001

a b s t r a c t

Insect pests infesting six stored botanicals were sampled monthly in two Egyptian warehouses over 12months. The plants sampled were; roselle (Hibiscus sabdariffa), mogat (Glossostemon bruguieri), coriander(Coriandrum sativum), anise (Pimpinella anisum), chamomile (Matricaria chamomilla) and marjoram(Origanum majorana). The warehouses were located in northern Egypt in Mansoura and Bilqas. Lasio-derma serricorne and Stegobium paniceum were the most common insect pests in warehouses. Thebeetles Tribolium castaneum, Tribolium confusum, Trogoderma granarium and Cryptolestes ferrugineus hadlower and similar levels of infestation. The moths, Plodia interpunctella and Sitotroga cerealella, had thelowest infestation levels. There were small differences in infestation by the most common insects,L. serricorne and S. paniceum, among the six botanicals. In general, M. chamomilla and O. majorana had thelowest level of infestation. The other plants, H. sabdariffa and C. sativum had higher levels of infestation.The warehouse in Bilqas had higher populations than the warehouse in Mansoura. In Bilqas, thetemperature and relative humidity were slightly higher and the warehouse was older and had openwindows, factors that may have contributed to higher insect infestations.

Crown Copyright � 2010 Published by Elsevier Ltd. All rights reserved.

1. Introduction

Most research in stored-product protection focuses on cereals,as cereals are the most common commodity in storage, and thereare several insects and moulds that can reduce the quality andquantity of cereals in storage. However, all agricultural productsare stored for some time before consumption, and they can alsobe infested by insects and moulds in storage. Dried plants, roots,stems, leaves, seeds and flowers, have been used from the dawnof history for medicine and as spices for cooking (Craker, 2007).Recently, there has been an increase in the popularity of herbalremedies in North America and Europe. However, there isa growing need to understand the problems in storing botanicals.Botanicals are also attacked in storage by several insects. Themost common insects found in botanicals are Lasioderma serri-corne (F.) and Stegobium paniceum (L.) (Lefkovitch, 1967; Tawfiket al., 1984; Awadallah et al., 1990; Arbogast et al., 2002). Theseinsects are even able to develop on Acorus calamus L. rhizomes(Srivastava and Saxena, 1975), a plant that has essential oils thatare toxic to several stored-product insects (El-Nahal et al., 1989).There are a number of other insects found in stored botanicals

: þ1 204 983 4604.

010 Published by Elsevier Ltd. All

(Araecerus fasciculatus (Degeer), Tenebroides mauritanicus (L.),Tribolium castaneum (Herbst), Oryzaephilus mercator (Fauvel),Ahasverus advena (Waltl), Thaneroclerus buqueti Lefebvre andThorictodes heydeni (Reitter) (Abraham, 1975; Tawfik et al., 1984;Basak, 1991)). There have been several studies examining insectsin stored botanicals worldwide; Egypt (Kamel, 1958; El-Halfawy,1977), India (Abraham, 1975; Basak, 1991), Germany (Schmidt andGraebner, 1987), Korea (Toh, 1998) and USA (Arbogast et al.,2002).

Egypt is a major producer of botanicals. Each year, Egypt exports7000 to 15,000 t of botanicals (Lange and Mladenova, 1997). Theseplants are grown in southern Egypt, harvested from June to August,dried, then placed in 50 kg jute bags and stored for 2–10 months inwarehouses before being shipped to processing facilities in Egypt,Europe and USA. As in other countries, the two most commoninsects in botanicals in Egyptian warehouses are L. serricorne and S.paniceum (Tawfik et al., 1984; Awadallah et al., 1990). These andother stored-product insects cause significant damage duringstorage. To reduce the damage, bag stacks are covered with a tarpand fumigated with phosphine derived from aluminum phosphide(Bond, 1984) once or twice during storage. However, there is anincreased demand from customers in Europe and USA for organicproduction, which does not allow the use of phosphine. Yet thereare very low tolerances for insect damage, live insects and insectfragments in stored botanicals.

rights reserved.

A.Y. Abdelghany et al. / Journal of Stored Products Research 46 (2010) 93–9794

Previous studies on insects in botanicals have described life-histories and losses caused by insect infestation during storage.Previous studies in Egypt have listed the insect pests and theirnatural enemies. The goal of this study was to sample quantitativelya wide variety of plants in storage throughout an entire year.

2. Material and methods

2.1. Warehouses

Sampling was conducted at two warehouses in the Delta regionof Egypt; Mansoura (31�00’N, 31�19’E) and Bilqas (31�31’N, 31�21’E)containing stored botanicals. The warehouse in Bilqas was older,had wooden floors and it was open to the environment; its area wasapproximately 240 m2. The warehouse in Mansoura had woodenfloors and was sealed from the outside environment with walls andwindows; its area was approximately 120 m2. Samples were takenonce a month from June 2005 to May 2006. Six dried plants weresampled; Hibiscus sabdariffa L. (roselle) fruits, Glossostemon bru-guieri (Dsef.) (mogat) roots, Coriandrum sativum L. (coriander)seeds, Pimpinella anisum L. (anise) seeds, Matricaria chamomillaL. (chamomile) flowers and Origanum majorana L. (marjoram)leaves. A sample of approximately 2.5 kg of each botanical wastaken in five locations in each warehouse (north, south, east, westand center).

2.2. Insect sampling

The plants are harvested from April to June each year. Farmers dryand store plants for a few weeks to a few months before delivering toa warehouse in June, July or August. Samples of plants were takenwith a spear sampler, then placed in paper bags and transferred tothe laboratory. In the laboratory, the samples were sieved twice with1.40 mm and 0.850 mm openings and any adult insects identifiedand counted. To estimate the immature stages, the plant materialwas held at 30�1 �C, and sieved again after 2, 4 and 6 weeks and anyadult insects identified and counted. The total number of insects,both adults and immatures, in the sample was used in the dataanalysis. In general, there were more adults than immatures in thesamples. Air temperature and relative humidity (r.h.) data were fromStevenson Screens at weather stations at Bilqas and Mansoura,maintained by the weather service of Dakhalia region. On the day ofsampling, warehouse air temperature was recorded each hour from12 midnight for 24 h. Average temperatures are reported. Sampleswere taken at approximately 12 noon.

2.3. Data analysis

We used a one-way ANOVA to compare the total number of allinsect species from all plants during the entire year from twowarehouses in Bilqas and Mansoura, Egypt, and a two-way ANOVAto compare the total number of L. serricorne and S. paniceumcollected during the entire year from six different plants in twowarehouses in Bilqas and Mansoura, Egypt. All means are reportedwith standard error of the mean (SEM).

3. Results

3.1. Seasonal abundance of insects

All insect species were present throughout the year on all plantmaterial, in both warehouses (Fig. 1). There was a low level ofinfestation at the beginning of the sampling. This increased grad-ually and reached the maximum in July and August in Mansouraand in September and October in Bilqas. In Bilqas, the mean total

number of all eight insect species collected during the twelvemonths of study was 2.0� 0.04 insects/kg. This was a significantlyhigher level of infestation than in Mansoura, which had1.7� 0.03 insects/kg (one-way ANOVA, F1,58¼ 200.3, P< 0.001).

3.2. Insects pests

In Mansoura, L. serricorne was the most common insect found,followed by S. paniceum. In Bilqas, the L. serricorne and S. paniceumwere more abundant than other species (Table 1). Other beetles,T. castaneum, Tribolium confusum du Val, Trogoderma granariumEverts and Cryptolestes ferrugineus (Stephens) occurred at the rateof approximately 0.24 individuals/kg in Bilqas and 0.12 individuals/kg in Mansoura. The two moths, Plodia interpunctella (Hubner) andSitotroga cerealella (Olivier) were the least common insect pestsfound in Bilqas.

3.3. Botanicals

There were small differences in infestation by the most commoninsects, L. serricorne and S. paniceum, among the six botanicals. Ingeneral, M. chamomilla and O. majorana had the lowest levels ofinfestation. Of the other plants, H. sabdariffa and C. sativum, hadhigher levels of infestation (Table 2).

3.4. Temperature and humidity

Temperature in both warehouses increased gradually froma minimum of approximately 13 �C in January, to a maximum of26–29 �C in September. Bilqas was warmer than Mansoura; theaverage annual temperature in Bilqas was 21.4�1.4 and in Man-soura it was 18.8� 1.2 (paired t-test, P< 0.001, t¼ 7.07) (Fig. 2B).The r.h. was relatively constant through the year and was between55% and 75%. Bilqas was more humid than Mansoura; averageannual r.h. in Bilqas was 69.1�1.3% and in Mansoura it was66.1�1.5% (paired t-test, P¼ 0.002, t¼�3.95) (Fig. 2A). For allinsect species in both Mansoura and Bilqas, there was a positivecorrelation between the average air temperature at the weatherstation and the insects found in botanicals in the warehouse(Table 3). In general, there was no correlation between r.h. and thenumber of insects. However, C. ferrugineus and P. interpunctella inMansoura showed a significant positive correlation between r.h.and the number of insects (Table 4).

4. Discussion

The insect population levels were different at the two locations.There are several possible reasons for these differences. Thewarehouse in Bilqas was older and was open to the environmentcompared to the newer, closed warehouse in Mansoura. Also, thewarehouse at Bilqas had more debris and broken bags on the floorthan in Mansoura. Sanitation can play an important role in reducingpest populations and in preventing outbreaks (Imholte andImholte-Tauscher, 1999). Initial populations may have been thesame in the two locations, but the poorer sanitation conditions inBilqas may have allowed for the much higher populations seen inSeptember and October. Finally, the temperatures and r.h. wereslightly higher in Bilqas, which would increase the rate of devel-opment and survival (Birch, 1953; Howe, 1957; Lefkovitch, 1967),and would lead to higher insect populations. Other possible reasonsfor the differences are the number of fumigations, the effectivenessof fumigations, use of contact insecticides and duration of storage.Both warehouses fumigated their stocks with phosphine, but wedid not examine the efficacy of fumigations. Further studies would

Fig. 1. Seasonal variation in the stored-product insects (individuals/kg) found in all six botanicals in two warehouses in Egypt.

A.Y. Abdelghany et al. / Journal of Stored Products Research 46 (2010) 93–97 95

be required to determine which of these factors are responsible forhigher insect populations in Bilqas.

The population dynamics and ecology are important in under-standing how and when it is best to control insect populations. Inthis study, insect populations rose in the late summer, peaking inJuly in Mansoura and in September in Bilqas. Howe (1957) esti-mates that there would be four to seven generations of L. serricornein Egypt. Declines in temperature were correlated with declines inpopulation densities. Although these insects can survive for

prolonged periods at 15–20 �C (Fields, 1992), temperatures seen inthe warehouses from November until May would cause femaleadults to stop laying eggs and completion of development to adultwould take months (Howe, 1957; Lefkovitch, 1967).

Similar trends are seen in other stored-product populations.Lasioderma serricorne had one or two population peaks in noodlefactories in Japan (Suezawa et al., 1987). Pscoids found in a tobaccoprocessing plant (Mashaya, 1999) mostly showed one and occa-sionally two peaks of population density each year. In an Italian

Table 1Total of all insects from all plants during an entire year from warehouses in Bilqas orMansoura, Egypt.

Insects Mean� SEM (individuals/kg)

Bilqas Bilqas

L. serricorne 0.31� 0.01a 0.23� 0.01aS. paniceum 0.27� 0.01ab 0.17� 0.007bT. castaneum 0.24� 0.01bc 0.13� 0.005cT. confusum 0.26� 0.009bc 0.13� 0.005cdT. granarium 0.23� 0.008bc 0.12� 0.005cdeC. ferrugineus 0.23� 0.007c 0.11� 0.002cdeP. interpunctella 0.19� 0.008d 0.11� 0.004cdeS. cerealella 0.18� 0.006d 0.10� 0.003e

For a given location, means that are significantly different are followed by differentletters, (Tukey’s test, P� 0.05).For Bilqas warehouse, one-way ANOVA, F7,232¼ 18.87, P< 0.001.For Mansoura warehouse, one-way ANOVA, F7,232¼ 40.49, P< 0.001.

Fig. 2. A. Monthly average daily air r.h. at weather stations in Bilqas and Mansoura,Egypt. B. Monthly average daily air temperature at Bilqas and Mansoura weatherstations, Egypt. Unconnected points are air temperatures taken in warehouses on daysof sampling.

A.Y. Abdelghany et al. / Journal of Stored Products Research 46 (2010) 93–9796

feed mill, there were large seasonal variations in insect populationsin the six insect species monitored but again there was a singlepeak in the population (Trematerra and Sciarretta, 2004). Similartrends are seen in grain, populations increase in the fall, there isa single peak in the population and populations decline again in thelate fall (Hagstrum, 1987; Sinha, 1987; Buchelos and Katopodis,1995). In grain residues in elevators in Kansas, USA, insect pop-ulations either had a single peak or two distinct peaks in populationdensity (Arthur et al., 2006).

There are many factors that affect the seasonal populationdensity of stored-product insects; immigration, emigration,temperature, humidity, food quality, longevity and survival, toname but a few (Birch, 1953; Lefkovitch, 1967; Price, 1997; Rees,2004). Control measures, such as fumigation, radically reducepopulations, and may alter the age structure of the population byleaving only the resistant stages such as eggs or pupae (Bond, 1984).The type of sampling can also affect estimates of populationdensity. Traps are often used to monitor populations (Suezawaet al., 1987; Mashaya, 1999; Hagstrum, 2000; Trematerra andSciarretta, 2004), but they are affected by many factors that areunrelated to population density, the main one being temperature(White et al., 1990). Direct sampling of the grain or product(Hagstrum, 1987; Sinha, 1987; Buchelos and Katopodis, 1995;Arthur et al., 2006, this study) is unaffected by temperature andgives an absolute density of the insect population in the storedproduct. However, clumped distribution and low numbers cancause difficulties in using this method.

There were only small differences in infestation between the sixdifferent botanicals. These plants were from widely different plantfamilies and also different parts of the plant were held in storage;seed, flower, fruit and leaves. In contrast, several stored-productbeetles survive better on certain cereals than others, and all thesehosts are relatively homogenous since all are seeds of the family

Table 2Total of L. serricorne and S. paniceum collected during an entire year from sixdifferent plants in warehouses, in Bilqas and Mansoura, Egypt.

Plants Mean� SEM (individuals/kg)

Bilqas Mansoura

L. serricorne S. paniceum L. serricorne S. paniceum

H. sabdariffa 2.26� 0.23ab 2.00� 0.20a 1.96� 0.06a 1.36� 0.13aC. sativum 2.26� 0.10a 1.76� 0.13ab 1.50� 0.16b 1.00� 0.13abG. bruguieri 1.76� 0.16abcd 1.70� 0.03ab 1.40� 0.13bc 1.06� 0.06abM. chamomilla 1.56� 0.16 cd 1.33� 0.10b 1.43� 0.23bc 0.83� 0.13bP. anisum 2.00� 0.10abc 1.60� 0.13ab 1.23� 0.06bc 1.10� 0.10abO. majorana 1.46� 0.10d 1.56� 0.16ab 1.10� 0.06c 1.10� 0.06ab

For Bilqas (two-way ANOVA, plant: F5,192¼ 3.122, P¼ 0.010; insect: F7,192¼ 22.353,P< 0.001) for Mansoura (two-way ANOVA, plant: F5,192¼ 3.021, P¼ 0.012; insect:F7,192¼ 52.60, P< 0.001).

Gramineae. For example, Cryptelestes spp. and Rhyzoperthadominica (F.) were found in greater densities in wheat than maize,whereas Oryzaephilus surinamenis (L.) was found mostly on oats(Storey et al., 1983). Lasioderma serricorne has one of the greatestranges of hosts of all stored-product insects (Howe, 1957), and thismay be one reason that few differences in infestation were seenbetween the different plants in our study.

Other studies have shown differences in insect infestationbetween botanicals. In Indian warehouses, L. serricorne andS. paniceum are the most common insects found on ginger,turmeric, quince and nagkesar, but these insects were absent fromchicory, nutmeg, vanilla, cinnamon and pepper (Abraham, 1975;Basak, 1991). In German warehouses, S. paniceum was found on 10out of 15 products (Schmidt and Graebner, 1987). In Korea, 12 out of15 dried herbs were found infested with S. paniceum (Toh, 1998). Ina laboratory study, L. serricorne survived better on coriander flourthan fennel, caraway or flour (Magd El-Din, 2003).

Currently, managers of these two Egyptian warehouses fumigatein December and January, when the populations are at their lowestlevel and the warehouses are cool. One advantage of this scheduleis that the quantity of botanicals to be fumigated is much less thanin the fall. Botanicals arriving in the warehouse in June and July mayalready have insect infestations, and this may be the main reasonpopulations are so high in August and September. Controlling thesepopulations early in June and July could prevent the high pop-ulations seen in August and September. However, if the mainreasons for the peak in August and September are insects attackingthe botanicals once they arrive in the warehouse, and these insectsare immigrating from outside the warehouse, then fumigating

Table 3Regression between insects and temperature.

Insects Mansoura Bilqas

R2 Slope Intercept P R2 Slope Intercept P

L. serricorne 0.70 23.97 13.06 0.0006 0.67 21.45 14.75 0.001S. paniceum 0.37 23.08 14.82 0.03 0.49 18.91 16.24 0.01T. castaneum 0.43 37.05 13.71 0.01 0.46 21.76 16.01 0.01T. confusum 0.39 59.32 10.78 0.02 0.38 18.42 16.72 0.03T. granarium 0.63 89.30 7.08 0.001 0.57 24.77 15.53 0.004C. ferrugineus 0.48 89.44 8.10 0.01 0.50 27.87 14.68 0.01P. interpunctella 0.58 92.87 8.00 0.003 0.56 48.46 11.87 0.004S. cerealella 0.358 80.23 10.14 0.03 0.33 32.28 15.55 0.04

Table 4Regression between insects and relative humidity.a

Insects Mansoura

R2 Slope Intercept P

C. ferrugineus 0.34 83.33 59.00 0.04P. interpunctella 0.52 97.23 57.73 0.008

a Only insects with significant correlations are listed.

A.Y. Abdelghany et al. / Journal of Stored Products Research 46 (2010) 93–97 97

stocks in June and July would have little effect on the peak. Fumi-gation would be better later in the summer, just before the begin-ning of cooler temperatures and fewer immigrating insects. Furtherresearch is required to determine which would be the best strategy.Regardless, fumigations would be more effective at the highertemperatures in June through September (Bond, 2004).

Fumigation would not prevent reinfestation. Increased sanita-tion would reduce beetles within the warehouse and should lead tolower populations within the botanicals. Changing from jute toa more impermeable bag material would limit the spread ofinfestation. This study shows that all botanicals have similar levelsof infestation, so that all plants would need to be treated. Since L.serricorne is the most common insect found, and product samplingis labour intensive, pheromone trapping could be an effective toolfor monitoring the effectiveness of a control program. Pheromonetraps catch fewer insects at lower temperatures, so this methodmay, however, be less effective from December to February whentemperatures are below 16 �C.

Acknowledgement

We would like to thank the Egyptian government for theirsupport of Ahmed Abdelghany, and Marjorie Smith for her statis-tical advice (AAFC, Winnipeg, Canada).

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