Altered olfactory receptor neuron responsiveness in rare Ostrinia nubilalis males attracted to the O. furnacalis pheromone blend

Three percent of E-strain Ostrinia nubilalis males fly upwind in response to the Ostrinia furnacalis pheromone blend [a 40:60 ratio of (E)-12-tetradecenyl acetate to (Z)-12-tetradecenyl acetate (E12-14:OAc to Z12-14:OAc)], in addition to their own pheromone blend [a 99:1 ratio of (E)-11-tetradecenyl acetate to (Z)-11-tetradecenyl acetate) (E11-14:OAc to Z11-14:OAc)]. We assessed the olfactory receptor neuron (ORN) responses of these behaviorally "rare" males versus those of normal males. For the three ORNs housed within each sensillum, we tested responsiveness to Z12-14:OAc, E12-14:OAc, Z11-14:OAc, E11-14:OAc, and the behavioral antagonist (Z)-9-tetradecenyl acetate (Z9-14:OAc). Z11-14:OAc, E11-14:OAc, and Z9-14:OAc stimulated ORNs exhibiting distinct small, large, and medium spike sizes, respectively. For rare and normal males, both Z12-14:OAc and E12-14:OAc usually elicited responses from the largest-spiking ORN. In many ORNs of normal males, Z12-14:OAc or E12-14:OAc stimulated the smaller-spiking ORN that is responsive to Z11-14:OAc. In rare males, detectable ORN responses from the smaller-spiking ORN in response to Z12- and E12-14:OAc were virtually non-existent. These differences in ORN tuning in rare males will tend to create an ORN firing ratio between the large- and small-spiking ORNs in response to the O. furnacalis blend that is similar to that elicited by the O. nubilalis blend.     

Sex pheromone of the butterbur borer, Ostrinia zaguliaevi

The sex pheromone blend of the butterbur borer, Ostrinia zaguliaevi (Lepidoptera: Pyralidae) was analyzed by means of gas chromatography-electroantennographic detection (GC-EAD), GC-mass spectrometry and a series of wind-tunnel bioassays. Four EAD-active compounds were detected in the female sex pheromone gland extract, and these were identified as tetradecyl acetate (14:OAc), (Z)-9-tetradecenyl acetate (Z9-14:OAc), (E)-11-tetradecenyl acetate (E11-14:OAc) and (Z)-11-tetradecenyl acetate (Z11-14:OAc). The average amounts ± s.d. of the four compounds in a single sex pheromone gland were 7.9±3.7 ng, 10.1±3.2 ng, 1.1±0.5 ng and 11.6±5.1 ng, respectively. In a wind-tunnel bioassay, the ternary blend of Z9-, E11- and Z11-14:OAc at a ratio found in the sex pheromone gland (45:5:50) elicited the same behavioral responses from the males as did virgin females and pheromone gland extract. Removal of any single compound from the ternary blend significantly diminished the pheromonal activity, whereas addition of 14:OAc to the ternary blend had no effect on the males' behavioral responses. Therefore, it was concluded that the sex pheromone blend of O. zaguliaevi is composed of Z9-14:OAc, E11-14:OAc and Z11-14:OAc at a ratio of 45:5:50.     

Identification of the sex pheromone of Ostrinia palustralis

By means of gas chromatography with electroantennographic detection, gas chromatography-mass spectrometry and a series of bioassays, (E)-11-tetradecenyl acetate (E11-14:OAc) and (Z)-11-tetradecenyl acetate (Z11-14:OAc) at a ratio of 99:1 were identified as female sex pheromone components of Ostrinia palustralis. The average amounts of E11- 14:OAc and Z11-14:OAc in a single sex pheromone gland were 37.2±24.4 ng and 0.3±0.2 ng, respectively. In a wind-tunnel bioassay, the binary blend of E11- and Z11-14:OAc elicited the same male behavioral responses as did virgin females.     

Evidence of olfactory antagonistic imposition as a facilitator of evolutionary shifts in pheromone blend usage in Ostrinia spp. (Lepidoptera: Crambidae)

Olfactory receptor neuron (ORN) response was measured to assess why some males ("rare males") of the Asian corn borer (ACB), Ostrinia furnacalis, have a broad behavioral response to fly upwind to both the ACB and the European corn borer (ECB), Ostrinia nubilalis, pheromone blends. We performed single-sensillum electrophysiological recordings on ACB males that had been behaviorally assessed for upwind flight response to the ACB blend [60:40 (Z)-12-tetradecenyl acetate (Z12-14:OAc) to (E)-12-tetradecenyl acetate (E12-14:OAc)], as well as to ECB (Z-strain) and ECB (E-strain) blends [3:97 and 99:1 (Z)-11-tetradecenyl acetate (Z11-14:OAc) to (E)-11-tetradecenyl acetate (E11-14:OAc)]. Sensilla from all types of males had large- and small-spike-sized ORNs responding strongly to Z12- or E12-14:OAc, but weakly to Z11- and E11-14:OAc. In the majority of males ("normal males") that flew upwind only to the ACB blend, Z11-14:OAc elicited responses in an intermediate spike-sized ORN associated with behavioral antagonism that is mainly tuned to (Z)-9-tetradecenyl acetate (Z9-14:OAc). In the rare-type ACB males that flew to both the ACB and ECB pheromone blends, Z11-14:OAc did not stimulate this ORN. Increased responsiveness to ancestral pheromone components by ORNs associated with behavioral antagonism could be instrumental in reproductive character displacement, or in reinforcement and reproductive isolation during speciation by helping to increase assortative mating between males and females in derived populations that use novel sex pheromone blends.     

Sex pheromone of Etiella behrii, a pod borer of soybean in Indonesia: identification and field attraction

Four EAG-active components were detected in GC-EAG analyses of hexane extracts from virgin Etiella behrii (Zeller) (Lepidoptera: Pyralidae) females. These components were identified as dodecyl acetate (12:Ac), (E)-9-dodecenyl acetate (E9-12:Ac), either (Z)-9- or (E)-11-tetradecenyl acetate (Z9- or E11-14:Ac), and (Z)-11-tetradecenyl acetate (Z11-14:Ac) by comparison of retention indices on both nonpolar and polar GC columns. The available extract was insufficient for further GC-MS or other chemical analyses (<0.2 ng/female). In field tests carried out in East Java, a 10:90 mixture of E9-12:Ac and Z11-14:Ac showed attractiveness to male moths and addition of 12:Ac and/or E11-14:Ac significantly increased the trap catches while addition of Z9-14:Ac showed no significant effect. Maximum attraction was obtained with 5.35 or 10.7 g/rubber septum of a mixture of E9-12:Ac, Z11-14:Ac, 12:Ac and E11-14:Ac at the ratio of 10:90:0.7:6.3, respectively. The role of pheromone blends in species discrimination between E. behrii and the related E. zinckenella (Treitschke) is discussed.     

大螟雄蛾触角对性外激素及其类似物的电生理反应

电生理试验显示出:大螟Sesamia inferens(Walker)雄蛾触角对Z11-16:Ac(A)oZ11-16:OH(B)激起强烈的触角电位(EAG)反应,Z11-16:ALD?,E11-16:Ac(D),Z9-14:Ac(E)和Z7-12:Ac(F)四种化合物也能激起较强的EAG反应;A化合物的刺激阈比B高,此两种化合物5:5比例时,激起的反应最强烈。电生理的试验推测,雄蛾触角对A和B两种化合物的感受,是通过触角内两种不同类型的嗅觉感受器。A和B不同比例混合后,田间诱蛾试验证明:在9:1,8:2,5:5的比例时,其诱蛾量远远超过单独的A化合物;E加入A与B的混合物中,起抑制诱蛾的效果,上述试验表明,Z11-16:OH是大螟的性诱剂。     

Response specificity of male olfactory receptor neurones for the major and minor components of a female pheromone blend

Abstract The sex attractant of the female redbanded leafroller moth, Argyrotaenia velutinana (Walker), is a blend of seven compounds. Specialized olfactory receptor neurones had been found for only two of the compounds, (Z)-11-tetradecenyl acetate (Z11-14:Ac) and (E)-11-tetradecenyl acetate (E11–14:Ac). These receptor neurones were always found in pairs within the long trichoid sensilla, which are the most abundant multi-pored sensilla on the male antenna. A systematic survey of all regions of the male antenna with standard extracellular recording techniques was undertaken to find receptor neurones responsive to the remaining five minor components of the female pheromone. Of the 113 long trichoid sensilla sampled, all contained two receptor neurones, one specialized for Z11–14:Ac and a second specialized for Ell –14:Ac. A comparable number of recordings were then obtained from the less abundant classes of multi-pored sensilla. Two new receptor neurone types were found, responsive to the minor pheromone components (E)-9-dodecenyl acetate (E9-12:Ac) and (Z)-9-dodecenyl acetate (Z9-12:Ac). Scanning electron micrographs indicated that these recordings were obtained from shorter, narrower trichoid sensilla. The majority of these sensilla appeared to contain three neurones capable of spontaneous action potential production. In each sensillum, only one receptor neurone appeared to respond to stimulation with a minor component of the female blend. The remaining two neurones did not respond to the chemical stimuli evaluated.     

Notes on New Zealand mammals 10. Effect of observer position on group size and level of aggression of mating brown hares

Abstract

The sex pheromones of two endemic New Zealand tortricid species were investigated. Females of the first species, Cnephasia jactatana, a pest of kiwifruit, were found to produce (Z)-11-tetradecenyl acetate (Z11-14:OAc) in their sex pheromone gland. When tested in the field, this compound on its own elicited significant catches of male C. jactatana. However, the addition of even very small amounts (0.3%) of the geometric isomer, (E)-11-tetradecenyl acetate (E11-14:OAc), to Z 11-14:OAc virtually suppressed trap catches. The sex pheromone of the second species, Merophyas leucaniana, was found to consist of a mixture of Z11-14:OAc, E11-14:OAc, tetradecyl acetate, and (Z)-11-tetradecenol. This species is morphologically very similar to the congeneric species, M. divulsana, the Australian lucerne leafroller. The sex pheromone of M. leucaniana is also very similar to that of the so-called “Z-type” of M. divulsana, which is now found in New Zealand. The pheromone components of C. jactatana and M. leucaniana are discussed in relation to those found in other tortricids.     

Relative attractiveness of incomplete and complete blends of synthetic pheromone to male obliquebanded leafroller (Lepidoptera: Tortricidae) moths in a flight tunnel and in apple orchards: implications for sex pheromone-mediated mating disruption of this species

The relative attractiveness of synthetic Choristoneura rosaceana (Harris) pheromone consisting of the major compound Z-11-tetradecenyl acetate (Z11-14:OAc), or the major compound plus one, two, or all three of the minor compounds E-11-tetradecenyl acetate (E11-14:OAc), Z-11-tetradecenol (Z11-14:OH), and Z-11-tetradecenal (Z11-14:Ald), in amounts similar to those in the natural pheromone, was compared in a flight tunnel and in apple orchards. In the flight tunnel, there was an increase in the proportion of moths that were activated when E11-14:OAc was combined with the main compound, but no detectible additional increases in response with the addition of Z11-14:OH and Z11-14:Ald. The time required for activation was twice as long when using the major compound than when using the major compound and one or more of the minor compounds. There was a trend of increase in the proportion of males initiating the take-off, lock-on, close-in, and touchdown phases of upwind flight when progressively more complete blends were used. The time required for the initiation of these behavioral phases was statistically similar for each treatment. The proportion of moths landing at the pheromone source was 10 times greater when using the complete blend than when using the main compound alone. Similar results were obtained in apple orchards, where the four-compound blend was 17-55 times more attractive than the main compound. The superior attractiveness of the four-compound pheromone compared with the major compound, or to the major compound plus the minor compound E11-14:OAc, suggests that it would be the most effective synthetic pheromone if competitive attraction were an important mechanism of disruption in this species.     

Reidentification of the female sex pheromone of the Indian meal moth, Plodia interpunctella: evidence for a four-component pheromone blend

Pheromone gland extracts from calling female Plodia interpunctella contained at least seven compounds that consistently elicited electroantennographic responses from male antennae upon gas chromatographic analysis. Three of these compounds were found to be the previously identified gland constituents, i.e., (Z,E)-9,12-tetradecadienyl acetate (Z9,E12-14:OAc), (Z,E)-9,12-tetradecadienal (Z9,E12-14:Ald) and (Z,E)-9,12-tetradecadienol (Z9,E12-14:OH). A fourth EAD-active compound was identified as (Z)-9-tetradecenyl acetate (Z9-14:OAc). The homologue (Z)-11-hexadecenyl acetate (Z11-16:OAc) was also identified in the extracts, but showed no EAD activity. The identity of all five compounds was confirmed by comparison of GC retention times and mass spectra with those of synthetic standards. In flight tunnel tests there were no significant differences in response of male P. interpunctella to the bait containing all four EAD-active compounds and the responses to female gland extacts. A behavioural assay of different two-compound blends in the flight tunnel showed that only addition of the corresponding aldehyde to the major pheromone component Z9,E12-14:OAc raised the male response. A subtractive assay, however, revealed that the exclusion of any of the compounds from the complete four-compound blend reduced its activity significantly. We thus conclude that the female-produced sex pheromone of P. interpunctella consists of at least four components, i.e., Z9,E12-14:OAc, Z9,E12-14:Ald, Z9,E12-14:OH and Z9-14:OAc.In a field trapping test performed in a storage facility, the four-component blend attracted significantly more males of P. interpunctella than traps baited with Z9,E12-14:OAc alone. In contrast, the highest number of Ephestia kuehniella males was found in the traps baited with this major component, suggesting that the secondary pheromone components contribute to the species specificity of the blend.     

Sulfhydryl-reagent inhibition of olfaction in a moth and effects of exposure to pheromone compounds or a pheromone analogue

The effects on olfaction of N-ethylmaleimide (NEM), a specificreagent of free sulfhydryl groups, were studied in the mothMamestra brassicae. The antennae of male M.brassicae bear twotypes of specialist receptor neurons involved in pheromone communication.One type is tuned to (Z)-11-hexadecenyl acetate (Z11-16:Ac),the main pheromone component; the second type is tuned to (Z)-9-tetradecenylacetate (Z9-14:Ac), an interspecific inhibitor not producedby the females of this species. Vapours of NEM irreversiblyinhibited the electro-antennographic (EAG) responses to Z11-16:Acand Z9-14:Ac. When Zll-16:Ac was applied before and during NEMtreatment, the responses to Z9-14:Ac were preserved and someprotection was observed in the responses to Zll-16:Ac. In return,Z9-14:Ac partially prevented the disappearance of responsesto Zll-16:Ac but not to Z9-14:Ac. A third compound, hexadecylacetate (16:Ac), found in the pheromone gland, but not detectedby the antennal receptors, did not prevent the inhibition causedby NEM.     

Evolution of noctuid pheromone binding proteins: identification of PBP in the black cutworm moth,Agrotis ipsilon

Male black cutworm moths (Agrotis ipsilon, Lepidoptera, Noctuoidea, Noctuidae), which are attracted by a three-component pheromone blend ((Z)-7-dodecenyl acetate, Z7-12:Ac; (Z)-9-tetradecenyl acetate, Z9-14:Ac; (Z)-11-hexadecenyl acetate, Z11-16:Ac), express diverse antennal pheromone binding proteins (PBPs). Two PBP isoforms (Aips-1 and Aips-2) that show 46% identity were cloned from antennal cDNA of male A. ipsilon. The protein Aips-1 displays a high degree of identity (70-95%) with PBPs of other noctuiids, but shows only 42-65% identity with the PBPs of more phylogenetically distant species. The other protein, Aips-2, represents a distinct group of PBP that includes proteins from Sphingidae and Yponomeutidae. These differences observed suggest that each of the two PBPs may be tuned to a specific pheromone ligand.     

A direct search approach to characterize the sex pheromone composition giving maximal male response

ABSTRACT. Response of male moths to sex pheromones differs markedly depending on blend composition. Natural selection can therefore be expected to favour males responding maximally to the blend produced by most females in the population. Thus it is relevant to quantify male response by the blend to which the males of a population respond maximally.
To do this, wind tunnel experiments were carried out where males were observed while responding to blends of synthetic compounds emitted from rubber septa. By stepwise improvement it was shown that the blend to which Yponomeuta cagnagellus responds maximally comprises (Z)11-tetradecenyl acetate (Z11-14Ac), ( E )11-tetradecenyl acetate (E11-14Ac), (Z)11-tetradecenol (Z11-140H), and tetradecyl acetate (14Ac) in the ratio 100(=50μg):1.5:20:45.
The innovate aspects of the method used to obtain this result, are the use of a direct search approach as an integral part of the experimental procedure and the use of a summary variable to characterize male response. These aspects are discussed using the data on the response of Y. cagnagellus males.     

豆荚螟中国种群性信息素鉴定

利用气相色谱一质谱联用技术（GC—MS）和气相色谱与风洞技术对交配期的豆荚螟（Etiella zinckenella）处女雌蛾的腺体提取物进行定性分析并侦测其中的活性组分,最终确定豆荚螟性信息素的有效组分为顺9一十四碳烯醇乙酸酯（z9—14AC）、顺11一十四碳烯醇乙酸酯（Z11—14AC）、十四碳烯醇乙酸酯（14AC）,三种成分在腺体中的比例分别1：3：1的存在。     

Antennal sensitivity to female sex pheromone compounds of Spodoptera frugiperda males (Lepidoptera: Noctuidae) and associated field behaviour

The fall armyworm (FAW), Spodoptera frugiperda (J. E. Smith) (Lepidoptera: Noctuidae), is one of the most important insect pests of corn, sorghum, rice, and grasses. The sex pheromone produced by S. frugiperda is composed of a mixture of esters. In this study, I determined the antennal responses of FAW males from 22 populations in Mexico to the components of the sex pheromone of this species: (Z)-7-dodecenyl acetate (Z7-12:OAc), (Z)-9-tetradecenyl acetate (Z9-14:OAc), and (Z)-11-hexadecenyl acetate (Z11-16:OAc) at four doses (0.01, 0.1, 1, and 10 μg). In addition, I evaluated the behavioural activity of single compounds in the field. The largest antennal responses of FAW males from all the sampled regions were elicited by Z9-14:OAc (all doses evaluated) and Z7-12:OAc (0.01, 0.1, and 1 μg). The male antennal responses evoked by Z11-16:OAc were smaller but similar to responses to Z7-12:OAc at lower doses (0.01 and 0.1 μg). In the field test, Z9-14:OAc produced significantly higher captures of S. frugiperda males compared to Z7-12:AOc and Z11-16:OAc. Antennal responses were related to captures obtained in the field, supported by the positive correlation of these variables [electroantennography (EAG) response and behaviour], with the exception of Z7-12:OAc, which elicited relatively large EAG responses but did not produce high captures of S. frugiperda. These results are of importance to agriculture, enabling the implementation of better methods of monitoring and controlling this pest, and support the suggestion that a mixture of Z9-14:OAc and Z7-12:OAc could represent an effective attractant for all evaluated populations of S. frugiperda in Mexico.     

Sex pheromone of the African armyworm <Emphasis Type="Italic">Spodoptera exempta</Emphasis> (Lepidoptera: Noctuidae): identification of components of the Okinawan population and formulation for population monitoring

In 2010, abrupt outbreaks of the African armyworm, Spodoptera exempta (Walker), occured on the Tarama, Iriomote and Kikai Islands in southwestern Japan. Analysis by gas chromatography-electroantennographic detection (GC-EAD) revealed two EAG-active compounds on male antenna in crude extract of virgin females. These compounds were identified as (9Z)-9-tetradecenyl acetate (Z9-14:Ac) and (9Z,12E)-9,12-tetradecadienyl acetate (Z9E12-14:Ac) in ca. 90:10 ratio by subsequent GC-MS analyses. (11Z)-11-Hexadecenyl acetate (Z11-16:Ac), which had previously been identified as a third component in the Kenyan population, was not detected. Binary blends of Z9-14:Ac and Z9E12-14:Ac at ratios between 99:1 and 90:10 showed a potent attractiveness in the field, superior to that of virgin females and comparable to that of the three-component formulation determined in Kenya. For the population survey, a 98:2 blend was used. In Tarama, only a few moths of S. exempta were captured with a light trap during the night when more than 600 males were captured with synthetic sex pheromone; more S. exempta captures with a light trap had been reported than with sex-pheromone traps in Kenya. This indicates that the Okinawan population has different properties from the Kenyan population in pheromone composition and behavioral response to light.     

Sex pheromone components of the leafroller moth Pandemis cerasana

Female-tip washings of the leafroller moth, Pandemis cerasana, were found to contain (E)-11-tetradecenyl acetate, (Z)-11-tetradecenyl acetate, (E)-11-tetradecenol, (Z)-11-tetradecenol and tetradecyl acetate, based on chemical analysis and electroantennogram tests. The relative amounts of these compounds in the gland were ca. 64:21:10:3:2 in the order named. Only (E)-11 and (Z)-11-tetradecenyl acetate were required for attraction of males to trap dispensed in the ratio 3:1, respectively.     

小地老虎中国种群性信息素的成分鉴定及田间试验（英文）

研究了小地老虎中国种群的性信息素组分,3日龄处女蛾单腺体性信息素提取物中性信息素的含量非常低。GC和GC-MS分析表明,小地老虎性信息素含有5种成分：顺-7-12碳乙酸酯（A）、顺-9-14碳乙酸酯（B）、顺-11-16碳乙酸酯（C）、顺-5-10碳乙酸酯（D）和顺-8-12碳乙酸酯（E）。它们的含量分别为：（0.245±0.098）、（0.080±0.031）、（0.089±0.03）、(0.085±0.031)和 (0.105±0.065)ng, 这5种物质的百分比分别为40.451±13.66、13.176±5.279、14.943±5.142、14.392±6.10和17.225±9.792,前3种物质的百分比为58.75±9.429、18.91±7.539和22.34±7.209。田间试验表明,性信息素单一组分均未诱到雄蛾,AB以3∶1的比例配成的诱芯对雄蛾有一定的引诱活性,一个诱捕器平均诱捕到2.6头。ABC组分以3∶1∶1的比例配成的诱芯对雄蛾引诱活性显著增强,一个诱捕器平均诱捕量为7.40头,是AB（3∶1）诱芯的2.8倍。诱芯中性信息素的含量对诱蛾活性有明显的影响,在剂量为200 µg时的平均诱捕量最高。     

Identification of the sex pheromone of Isoceras sibirica Alpheraky (Lepidoptera, Cossidae)

We discovered that extracts of the female sex pheromone gland of the carpenterworm moth Isoceras sibirica Alpheraky, a pest of Asparagus officinalis Linn., contained (Z)-7-tetradecen-1-ol (Z7-14:OH), (Z)-9-tetradecen-1-ol (Z9-14:OH), (Z)-7-tetradecenyl acetate (Z7-14:Ac), (Z)-9-tetradecenyl acetate (Z9-14:Ac), and (Z)-9-hexadecadecenyl acetate (Z9-16:Ac). The average levels of the chemicals in a single sex pheromone gland of a calling moth were (0.71 +/- 0.24) ng, (1.42 +/- 0.44) ng, (4.36 +/- 0.32) ng, (8.71 +/- 0.26) ng, and (0.82 +/- 0.38) ng, respectively. The electroantennography (EAG) analysis of these chemicals and their analogues demonstrated that Z9-14:Ac triggered significantly the male EAG response. Traps with rubber septa lure impregnated with Z9-14:Ac (500 microg/septum), Z7-14:Ac (250 microg/septum), and Z9-16:Ac (50 microg/septum) were more effective in catching male moths than traps with other baits or virgin females. Addition of Z7-14:OH and Z9-14:OH to rubber septa did not enhance the efficiency of the trap.     

Field trials of synthetic sex pheromone lures for the large cabbage-heart caterpillar moth, Crocidolomia pavonana (Lepidoptera: Crambidae) in Indonesia

The efficacy of synthetic female sex pheromone lures for Crocidolomia pavonana (Fabricius) (Lepidoptera: Crambidae) in the cabbage fields of Java and Bali, Indonesia, was investigated by varying the composition and dosage of the components. The lure containing a synthetic pheromone blend of (Z)-11-hexadecenyl acetate (Z11–16: Ac) and (Z)-9-tetradecenyl acetate (Z9–14: Ac) at a 10:1 ratio acquired significantly more male catches than single component lures and the control lure. Meanwhile, no attraction was observed when lures with 1:1 and 1:10 blends were tested. The composition of Z11–16: Ac and Z9–14: Ac at a ratio of 5, 10 and 20:1 attracted more males than the control lures. Dosage studies showed that 0.055 and 0.55 mg of a mixture of Z11–16: Ac and Z9–14: Ac (10:1 ratio) attracted more males than the control. These results are the first demonstration of the efficacy of synthetic pheromone for C. pavonana in field conditions. The present study suggests the feasibility of pheromone-based monitoring as a simple and low-cost technique for integrated pest management of this pest.