Sex biased intra-tree oviposition site selection and larval foraging behavior of a specialist herbivore

Heterogeneity in the quality of oviposition and feeding sites within plants can significantly influence the distribution and abundance of herbivorous insects, but remains poorly understood. Field surveys and a manipulative study were conducted to evaluate the influence of variation within the crown of black spruce, Picea mariana (Mill.) B.S.P. (Pinaceae), on adult oviposition and larval feeding behavior of yellowheaded spruce sawfly, Pikonema alaskensis Rohwer (Hymenoptera: Tenthredinidae). Most eggs were laid in the mid to lower crown of 1.5–2 m tall trees. However, most of the few eggs that were laid in the upper crown (i.e., whorl 2) were female. Fourth and fifth instars dispersed acropetally, from the mid and lower to the upper crown, causing high defoliation in the upper crown. Late-instar females were generally more abundant than males on the leader, the most apical shoot on a tree where eggs and early instars rarely occurred, strongly suggesting that more females than males disperse acropetally. This hypothesis was supported in a manipulative experiment, where only 15–20% of larvae in all-male broods, but almost three-quarters of larvae in mixed broods, dispersed to the upper crown. To our knowledge, this is only the second study to explicitly demonstrate preferential allocation of progeny sex through oviposition site selection by a herbivorous insect, and the first study to unambiguously demonstrate sex-biased dispersal by the juveniles of an insect whose adult females can fly. This study emphasizes the important role of intra-plant variation in shaping both oviposition site selection and the dispersal behavior of juvenile phytophagous insects within their hosts, and suggests that sex-biased foraging behaviors may be necessary for some insects to accommodate the respective needs of immature females and males within heterogeneous host plants.     

Influence of intra-tree variation in time of budburst of white spruce on herbivory and the behaviour and survivorship of Zeiraphera canadensis

Abstract.

• 1 Field studies were carried out to determine the effects of intra-tree variation in the time of budburst of white spruce, Picea glauca Moench (Voss.), on the behaviour and survivorship of, and herbivory by, the spruce bud moth, Zeiraphera canadensis Mutt. & Free.
• 2 There was significant variation in the time of budburst among whorls, shoots and buds. Budburst was acropetal, with buds in the interior of the lower crown bursting first and terminal buds on terminal shoots in the upper crown bursting last.
• 3 Bud moths laid the greatest proportion of their eggs in the middle of the crown and egg hatch was usually best synchronized to budburst in this region. Many eggs hatched before terminal buds on terminal shoots in the upper crown had burst and thus intra-tree variation in budburst decreased the probability that first-instar larvae would colonize the most important plant parts for growth.
• 4 However, many later instars dispersed upwards and outwards in the crown and colonized the late bursting buds in the upper crown. Such dispersal reduces the effectiveness of intra-plant variation in budburst to reduce herbivory and permits Z.canadensis to eat young nutritious buds for a longer period of time.
• 5 Intra-tree variance in the date of budburst was greater than that between trees but there were no consistent differences between the intra-tree variance of trees in half-sib families with high or low susceptibility to Z.canadensis. Differences between trees in herbivory, bud moth density and survivorship were not related to the amount of intra-tree variation in budburst.

     

Induced tree response to herbivory influences the foraging preference and performance of a conifer‐defoliating sawfly

Herbivory often alters the growth and development of woody plants and can thereby render hosts less susceptible to subsequent herbivores. We carried out field surveys and experiments to investigate how previous herbivory influences adult egg lay, larval feeding preference, and associated survival of the yellowheaded spruce sawfly, Pikonema alaskensis (Rohwer) (Hymenoptera: Tenthredinidae), within crowns of black spruce, Picea mariana (Mill.) Britton et al. (Pinaceae). Pikonema alaskensis females laid nearly twice as many eggs, and late‐instar larvae consumed nearly twice as much foliar biomass, on undefoliated vs. defoliated branches. The major factor driving the lower incidence of egg lay on defoliated branches was a significant reduction in the availability of preferred size classes of shoots. In general, adult females preferred medium‐sized shoots, whereas late instars preferred large shoots; previous defoliation generally decreased the availability of medium and large shoots. In a field bioassay, late‐instar survival generally declined with increasing defoliation, which again corresponded to significant reductions in shoot length on defoliated branches. The tendency of P. alaskensis to avoid shoots on severely defoliated branches is likely to increase the time lag between herbivory and the feedback effects of induction and could thus contribute to fluctuations in sawfly population density during outbreak. Our results differ from numerous past studies of closely related diprionid sawflies in conifers, most of which have reported weak or positive effects of previous herbivory on host susceptibility; we attribute the observed responses of spruce to the unusual sawfly feeding preference of P. alaskensis for developing conifer foliage.     

Dispersal and development of bagworm larvae (Lepidoptera: Psychidae) on three host plants

The present study investigated the development and dispersal of bagworm larvae, Thyridopteryx ephemeraeformis (Haworth) (Lepidoptera: Psychidae), on three hosts: arborvitae, Thuja occidentalis, blue spruce, Picea pungens, and juniper, Juniperus virginiana. The biomass of larvae was recorded at repeated intervals at 21 field sites. The relative quality of plants varied during larval development: early in the season, larvae were heavier on juniper and arborvitae than on blue spruce; late in the season, in contrast, larvae and pupae were lighter on juniper than on the two other plants. Larvae on blue spruce pupated later than those on arborvitae. In the laboratory, feeding rate and larval growth were not influenced by the density of larvae on the foliage of arborvitae up to a range which greatly exceeded that observed at any of our experimental sites. The incidence of dispersal among neonates emerging on arborvitae was high, as indicated by the high number of larvae captured on sticky traps, and the relatively low number of early instars on arborvitae foliage in relation to the reproductive output of females in the parental generation. The high rate of dispersal among early instars may be due to potential resource depletion over successive generations of bagworms.     

Influence of intra‐tree variation in phenology and oviposition site on the distribution and performance of Ennomos subsignaria on mature sycamore maple

Abstract 1. Field surveys and a manipulative experiment were conducted to examine the hypotheses that intra‐tree heterogeneity in natural enemy activity, foliar quality (independent of phenology), or phenology influence the intra‐tree distribution and performance of Ennomos subsignaria on mature sycamore maple, Acer pseudoplatanus. 2. Ennomos subsignaria intra‐tree distribution was distinctly clumped. Egg mass density was 85% higher on the lower bole than in the crown. Most early instars were found on lower crown proximal branches while most late instars and pupae were found on lower crown distal branches. This resulted in high levels of defoliation in the lower crown, especially on proximal branches. 3. No parasitoids were reared from eggs or late‐instar larvae and only one pupa was parasitised, suggesting that preference for the bole and lower crown was not a response to parasitism. Similarly, E. subsignaria performance was not influenced by variable foliage quality (independent of phenology) within the crown. However, sycamore maple phenology had a large influence on E. subsignaria survival. More than 90% of newly emerged larvae survived to the adult stage when they fed on foliage with three pairs of leaves expanded per bud, whereas survival on younger foliage was reduced by >45%. 4. The peak period of E. subsignaria egg hatch was approximately 2 weeks after the peak period of sycamore maple budburst, which occurred acropetally. Egg hatch was closely synchronised with the availability of most suitable leaves for insect development on proximal branches of the lower crown, the location where most larvae initiated feeding. 5. The results support the phenology hypothesis and suggest that intra‐tree variation in oviposition site and host phenological development influence the intra‐tree distribution and performance of this generalist herbivore.     

Sublethal effects of Bacillus thuringiensis on the spruce budworm, Choristoneura fumiferana

Exposing larvae of the spruce budworm, Choristoneura fumiferana (Clemens), to sublethal ( 50% lethal dose) levels of Bacillus thuringiensis subsp. kurstaki at various stages of their development significantly increased development time to the pupal stage and reduced pupal size and number of eggs laid per female, but did not affect the proportion of embryonated eggs. The changes in larval development time, pupal weight and fecundity depended on the larval stage that was treated. Exposure of fourth instars delayed larval development and reduced only male pupal weights with no effects on fecundity. Exposure of sixth instars delayed larval development to a lesser extent than exposure of fourth instars but had a pronounced effect on weight of both male and female pupae. The effect on pupal weight was sex dependent, as males tended to be more affected than females. The reduction in male pupal weight did not appear to influence fecundity, because the effect of exposure was explained by the change in female pupal weight. Effects on larval growth and pupal weight were proportional to the dose that was ingested during exposure, and were observed at doses as low as one-tenth of the LD₅₀. Ingestion of an LD₅₀ caused a 29 or 45% delay in development of, respectively, female or male larvae when exposed as fourth instars and a 30% reduction in female pupal weight when larvae were exposed as sixth instars.     

Temporal and spatial variations in microclimate influence the larval foraging behaviors and performance of a conifer-feeding sawfly

Herbivorous insects are often exposed to broad temporal and spatial variations in microclimate conditions within their host plants and have adapted a variety of behaviors, such as avoidance or basking, to either offset or benefit from such variation. Field experiments were carried out to investigate the influence of daily and intratree variations in microclimate on the behaviors (feeding, resting, dispersal, and hiding) and associated performance of late-instar larvae of the yellowheaded spruce sawfly, Pikonema alaskensis (Rohwer) (Hymenoptera: Tenthredinidae) within crowns of 1.25-1.5 m tall black spruce (Picea mariana [Miller] Britton Sterns Poggenburg); late instars feed on developing shoots of young spruce and are often exposed to microclimatic extremes with unknown effects on performance. Larvae fed diurnally from just after dawn (0800 h) until dusk (2000 h) and rested throughout the night, with brief periods of dispersal occurring in the morning and evening. Neither larval behavior nor abiotic conditions differed significantly between the upper and lower crowns of trees. Temperature, humidity, and solar insolation all explained >90% of variation in feeding; however, sunrise and sunset were the most likely cues influencing diurnal behavior. Most larvae (94%) fed on the bottom, shaded side of shoots, and field experiments indicated that this behavior is adaptive with respect to microclimate, probably reducing hygrothermal stress. Thus, behavioral adaptations by P. alaskensis to daily and within-shoot microclimatic variation may reduce the risk of hygrothermal stress during dispersal or feeding, while still allowing larvae to feed on the preferred and highly nutritious upper crown foliage of young spruce.     

Host tree influences on the dispersal of first instar gypsy moths, Lymantria dispar (L.)

Abstract. 1. In laboratory tests, first instar gypsy moths attempted dispersal more frequency when exposed to less acceptable foliage.
2. First instars from small eggs attempted dispersal less frequently than larvae from large eggs when exposed to foliage from highly acceptable or marginally acceptable hosts. Dispersal rates of larvae from medium sized eggs were intermediate.
3. These results (1–2) confirm and expand upon the findings of Capinera & Barbosa (1976).
4. In the field, data on the relative densities of larvae on different host species support the conclusion that the frequency of dispersal attempts is inversely related to host acceptability.
5. The implications of these findings for the population dynamics of the gypsy moth are discussed.     

Diamondback moth oviposition: effects of host plant and herbivory

The oviposition behaviour of Plutella xylostella L. (Lepidoptera: Plutellidae) on Chinese cabbage (Brassica rapa L. Pekinensis, cv. Wombok), canola (Brassica napus L. cv. Thunder TT), and cabbage (Brassica oleracea L. Capitata, cv. sugarloaf) (Brassicaceae) was studied in the laboratory. In no‐choice experiments moths laid most eggs on the stems and lower three leaves of cabbage plants, the lower three leaves of canola plants, but on the upper three leaves of Chinese cabbage plants. The effects of conspecific herbivore damage to foliage could be replicated by mechanical damage. When foliage was damaged, injured cabbage and canola plants were preferred for oviposition over intact conspecifics, whereas injured Chinese cabbage plants were less preferred than intact conspecifics. However, when root tissue was damaged, intact cabbage and canola plants were preferred over injured conspecifics, whereas moths did not discriminate between root‐damaged and intact Chinese cabbage plants. Injury to upper leaves significantly affected the intra‐plant distribution of eggs. In cabbage and canola plants, injury to leaf 6 significantly increased the number of eggs laid on this leaf, resulting in a significant decrease in the number of eggs laid on the lower foliage/stem of plants, whereas in Chinese cabbage plants it significantly decreased the number of eggs laid on leaf 6. Following oviposition on intact plants, neonate larvae established the vast majority of feeding sites on leaves 5–8 in all three host plants, indicating that larvae moved a considerable distance from preferred oviposition sites in cabbage and canola plants. The growth rate of neonates fed on leaf‐6 tissue was significantly greater than that of those fed on leaf‐1 tissue; >90% of larvae completed development when fed exclusively on leaf‐6 tissue but no larvae completed development when fed exclusively on leaf‐1 tissue. The study demonstrates the complex and unpredictable interactions between P. xylostella and its host plants and provides a basis from which we can begin to understand observed distributions of the pest in Brassica crops.     

Japanese oak silkmoth feeding preference for and performance on upper-crown and lower-crown leaves

We quantified differences in leaf traits between upper and lower crowns of a deciduous oak, Quercus acutissima, and examined feeding preference, consumption and performance of the Japanese oak silkmoth, Antheraea yamamai, for those leaves. Upper‐crown leaves had significantly smaller area, larger dry mass per area, greater thickness, lower water content, higher nitrogen content and a higher N/C ratio than lower‐crown leaves. When simultaneously offered upper‐crown and lower‐crown leaves, moth larvae consumed a significantly larger amount of the former. However, when fed with either upper‐crown or lower‐crown leaves (no choice), they consumed a significantly larger amount of the latter. Female larvae reared on upper‐crown leaves had a significantly smaller fresh weight, but attained a significantly larger pupal fresh and dry weight, with a significantly higher relative growth rate than those on lower‐crown leaves. Although, like female larvae, male larvae had a significantly smaller fresh weight when reared on upper‐crown leaves, they had a significantly larger value only for pupal dry weight. These results suggest that: (i) larvae ingest a greater amount of lower‐crown leaves to compensate for the lower nitrogen content of the foliage, resulting in having an excess of water because of the higher water content of the foliage; (ii) feeding preference for upper‐crown leaves accords with better performance (with respect to dry pupal weight and relative growth rate) on the foliage; (iii) better performance is explained by a higher nitrogen content and N/C ratio of the upper‐crown foliage; and (iv) the effects of leaf quality on performance differ between sexes.     

椰心叶甲实验种群的生物学特性观察

自然温度条件下 ,椰心叶甲Brontispalongissima (Gestro) 1年可发生 3代以上 ,世代重叠 ,主要以成虫越冬。成虫平均寿命 1 5 6d ,最长达 2 3 5d ,雌雄性比为 1∶1 ;雌雄虫一生均可交配多次。成虫产卵期长 ,产卵不规则 ,单雌平均产卵 1 1 9粒 ,最多可达 1 96粒。飞行磨测定的初步结果表明 ,雌虫飞行能力比雄虫强 ,2 4h未取食成虫最远飞行距离可达 40 0多m ;成虫和幼虫均具有负趋光性、假死性。成虫 3～ 5d不取食、高龄幼虫 7d不取食仍存活。幼虫经历 4～ 5龄 ,在温度或寄主不适宜条件下 ,可进入 6～ 7龄 ,或提前化蛹 ,从卵到成虫羽化大约需 3 6～ 61d。     

Observations on the life cycle and establishment ofCochylis atricapitana (Lep: Cochylidae), a moth used for biological control ofSenecio jacobaea in Australia

Larvae of the mothCochylis atricapitana (Stephens) are monophagous leaf, crown, stem or bud borers of ragwort,Senecio jacobaea L. (Asteraceae). In the present investigation, aspects of the life cycle ofC. atricapitana were determined. Moths ofC. atricapitana lay an average of 158 eggs/female with as many as 355 eggs being laid by a single female. The majority of eggs are laid individually along the primary and secondary veins on the underside of ragwort leaves. Egg incubation ranges from 4.2 days at 30°C to 14.4 days at 15°C. At a constant 23°C under a 16 hour photoperiod,C. atricapitana takes approximately 40 days to complete a generation. Caterpillars make their way to young, actively growing ragwort shoots or buds, and begin mining into the plant tissue, boring into the leaf, crown, stem or bud.C. atricapitana has five larval instars and enters diapause as a final instar larva. In southern Victoria, moths ofC. atricapitana fly from late September through to the beginning of February. Adults emerge after overwintering towards the end of spring or beginning of summer.C. atricapitana has established at two sites while larvae, or signs of damage have been observed at approximately 52% of release sites.      

Larval feeding behaviour affects the impact of staminate flower production on the suitability of balsam fir trees for spruce budworm

Laboratory rearing of spruce budworm, Choristoneura fumiferana, in conjunction with field rearing indicated that the feeding behaviour of the larvae, which is affected by the insect population density, significantly influenced the impact of balsam fir, Abies balsamea, staminate flowering on spruce budworm biology. At low budworm density, the production of pollen in the midcrown of host trees reduced the insect development time by 5 days without affecting pupal weight, fecundity and survival. However, at high budworm density, the small amount of current-year foliage produced by flowering branches forced old larvae (sixth instar) either to feed on 1-year-old foliage (backfeeding) or to move from the midcrown to the lower crown section where staminate flowers are absent and more current-year foliage is available. When old larvae fed on old foliage, they exhibited reduced pupal weight and fecundity without losing the advange in development time that they obtained from feeding on pollen during their early stages of development. On the other hand, when old larvae moved to the lower crown section, they avoided the negative effects of backfeeding but lost the advantage in development time that was gained from feeding on pollen. Results from this study indicated that the production of staminate flowers by balsam fir trees could have opposite effects on spruce budworm population dynamics depending upon the insect population density when flowering occurs.     

Reproductive Biology of Celatoria compressa (Diptera: Tachinidae), a Parasitoid of Diabrotica virgifera virgifera (Coleoptera: Chrysomelidae)

The tachinid Celatoria compressa, a parasitoid of adult Diabrotica species in North America has been studied as a candidate classical biological control agent for the western corn rootworm, Diabrotica virgifera virgifera, in Europe. Prior to its potential importation, a thorough understanding of the parasitoid's reproductive biology is essential, and is an important component in the evaluation of a species as a biological control agent. In this study it has been clarified that C. compressa belongs to a group of a few tachinid species characterised by having eggs that contain fully developed larvae which are laid directly into the host. After mating, the egg load of females increased steadily from day 1 to a maximum egg load on day 4. Thereafter eggs containing fully developed first instar larvae reached a maximum of 31 in 69 eggs. At the first day of larviposition, females laid on average only five eggs into multiple hosts, which is in contrast to the availability of 18 eggs containing fully developed first instars in the uterus per female at that time. During a mean female's larviposition period of 23 days, a total of 33 first instars were larviposited into the hosts, which is only half of the female's egg load. Lifetime fecundity of C. compressa was significantly correlated with longevity. However no relationship was found between body size and either lifetime fecundity or longevity. In this study, an inverse host density-dependent pattern of percent parasitism was shown for C. compressa under 24-h fixed-time laboratory conditions, reflecting a Holling type II response. The number of host parasitized per C. compressa female reached an upper limit of 10 hosts with an increasing host density, which can be explained by the long host handling time of C. compressa.     

蔷薇三节叶蜂的生物学及防治

在重庆为害月季的蔷薇三节叶蜂Arge przhevalskii Guss.一年发生7代;幼虫雌性6龄,雄性5龄,老熟后入浅层土中结茧,有预蛹期;温度和湿度是影响卵孵化及幼虫生长发育的重要因子;主要进行孤雌生殖;几十粒卵集中产于枝条上部的茎杆组织,排成两列,产卵的方位总是以枝条顶部为起点;初卵幼虫向顶爬行,1—3龄群集在枝条顶部的叶片为害,渐向下转移,4龄开始分散.20%速效菊酯通油35000～45000倍,25%多杀菊酯乳油4000—8000倍,80%敌敌畏乳油对3—4龄幼虫均有良好的防治效果.     

Spruce budworm growth,development and food utilization on young and old balsam fir trees

Laboratory rearing of spruce budworm, Choristoneura fumiferana, in conjunction with field rearing, gravimetric analyses, a transfer experiment, and foliage chemical analyses at six dates during the period of budworm feeding activity indicated that the age of balsam fir, Abies balsamea, trees (70-year-old mature trees or 30-year-old juvenile trees) affected tree suitability for the spruce budworm via the chemical profile of the foliage. Insects reared on old trees had greater survival and pupal weight, shorter development times, and caused more defoliation than those reared on young trees. Young trees were more suitable for the development of young larvae (instars 2–5), while old trees were more suitable for the development of older, sixth-instar larvae. These results were confirmed by the laboratory transfer experiment. Young larvae fed foliage from young trees had higher relative growth rates (RGR), digestibility (AD), and efficiency of conversion of ingested foliage (ECI) than those fed foliage from old trees. These differences appeared to be related to the high N:tannins ratio, and the high contents of P present in young trees during the development of the young larvae. Old larvae fed foliage from old trees had higher relative growth rates, relative consumption rates (RCR), and digestibility of the foliage than those fed foliage from young trees. The high digestibility of the foliage of old trees was compensated for by a lower efficiency of conversion of digested food (ECD), which in turn resulted in no significant effect of tree age on the efficiency of conversion of ingested foliage by old larvae. The low relative consumption rate of old larvae fed foliage from young trees appeared to be related to the low N:tannins ratio, and the high contents of bornyl acetate, terpinolene, and °-3-carene present in young trees during the budworm sixth instar. Variations in these compounds in relation to tree age may serve as mechanisms of balsam fir resistance to spruce budworm by reducing the feeding rate of sixth instar larvae.     

Life Cycle of the Mulberry Longicorn Beetle,Apriona germari Hope on an Artificial Diet

Mulberry longicorn beetle, Apriona germari, was reared on an artificial diet to investigate its life cycle. Average egg period was 17.95 days. Most larvae pupated at the 8th (33.3%) or 9th (33.3%) instars, but others emerged from the 7th to the 11th instars. The total duration of larval stage from the 1st to the end of the 9th and the 11th instars were 176.2 and 251.5 days, respectively. For terminatiopn of diapause and acceleration of pupation, low temperature treatment at 5°C for 60 days was highly effective. Average pupal period for female and male was 19.3 and 18.4 days, respectively. Average longevity of adults was similar with 40.5 days for female and 44.3 days for male. Mating had occurred from about 10 days after the adult emergence, and then a female adult laid one or two eggs per day. Average number of eggs oviposited by a female was approximately 47.7. Total life span of A. germari, when reared on an artificial diet in the laboratory, ranged broadly from 197.5 days to 331.5 days mainly depending on the larval period.     

Effects of nitrogen concentrations of Eucalyptus blakelyi foliage on the fecundity of Paropsis atomaria (Coleoptera: Chrysomelidae)

Summary The concentration of nitrogen in the foliage of Eucalyptus blakelyi strongly influenced both the total number of eggs laid and the rate of production of eggs by female Paropsis atomaria, a chrysomelid defoliator of Eucalyptus spp. Females feeding on foliage with low levels of nitrogen laid fewer eggs at a slower rate than those feeding on foliage with high levels of nitrogen. an increase from 1.5% to 4.0% of nitrogen in the foliage increased the total number of eggs laid by 500% and the rate of production of eggs by 400%. The rate at which females fed was not influenced by the concentration of nitrogen in their food. The viability of their eggs was negatively related with the concentration of nitrogen in the diet. Females feeding on foliage with low levels of nitrogen laid eggs which were significantly heavier than those of females on high nitrogen diets. Females on high nitrogen survived significantly longer than those on low nitrogen. In the field the concentration of nitrogen in the foliage of E. blakelyi occurs over a range shown to dramatically influence both the fecundity and rate of production of eggs of P. atomaria. Nitrogen may therefore play an important role in the population dynamics of this beetle.     

Protection of potato crop against Phthorimaea operculella (Zeller) infestation using frass extract of two noctuid insect pests under laboratory and storage simulation conditions

The oviposition deterrent effect of water extract of Spodoptera littoralis and Agrotis ipsilon larval frass on Phthorimaea operculella adult females was studied using two types of larval food “Natural host and Semi-artificial diet” under laboratory and storage simulation (semi-field) conditions. Extracted frass of fed larvae on semi-artificial diet showed complete oviposition deterrent effect at treatments with 4th, 5th and 6th instars of S. littoralis, also at treatments with 1st–3rd and 6th instars of A. ipsilon, while the same effect was observed when the larvae fed on castor oil leaves as a natural host only at treatment with frass extract of A. ipsilon 6th instar larvae. Presence of low amounts of phenols and flavonoids in water extract of A. ipsilon larval frass resulted in relatively more effect as oviposition deterrent to fertile adult females on treated oviposition sites, while the opposite effect was obtained in S. littoralis larval frass experiments. At semi-field experiments, the percentage reduction of laid eggs reached 100% after two?days at treatments with frass extracts of 4th and 5th S. littoralis larval instars and A. ipsilon 6th instar larvae fed on semi-artificial diet and/or castor oil leaves. Percentage reduction of laid eggs for untreated sacks reached 93.24 and 48.95% after 2 and 30?days, respectively, when placed between treated sacks, in comparison with the mean number of laid eggs for isolated control.